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NELSON MENOLLI JUNIOR
O gênero Pluteus no Brasil: revisão taxonômica
e contribuição à filogenia molecular
Tese apresentada ao Instituto de Botânica da
Secretaria do Meio Ambiente, como parte dos
requisitos exigidos para a obtenção do título de
DOUTOR em BIODIVERSIDADE VEGETAL
E MEIO AMBIENTE, na Área de Concentração
de Plantas Avasculares e Fungos em Análises
Ambientais.
SÃO PAULO
2013
NELSON MENOLLI JUNIOR
O gênero Pluteus no Brasil: revisão taxonômica
e contribuição à filogenia molecular
Tese apresentada ao Instituto de Botânica da
Secretaria do Meio Ambiente, como parte dos
requisitos exigidos para a obtenção do título de
DOUTOR em BIODIVERSIDADE VEGETAL
E MEIO AMBIENTE, na Área de Concentração
de Plantas Avasculares e Fungos em Análises
Ambientais.
ORIENTADORA: DRA. MARINA CAPELARI
Ficha Catalográfica elaborada pelo NÚCLEO DE BIBLIOTECA E MEMÓRIA
M547g
Menolli Junior, Nelson
O gênero Pluteus no Brasil: revisão taxonômica e contribuição à filogenia
molecular / Nelson Menolli Junior -- São Paulo, 2013.
273 p.il.
Tese (Doutorado) -- Instituto de Botânica da Secretaria de Estado do Meio
Ambiente, 2013
Bibliografia.
1. Basidiomicetos. 2. Agaricales. 3. Taxonomia. I. Título
CDU : 582.284
À minha mãe Alda Cristina Facion Menolli e ao
meu pai Nelson Menolli, base de todo meu ser e
responsáveis por cada pedacinho das minhas
conquistas.
AGRADECIMENTOS
Ao Instituto de Botânica e ao Programa de Pós-graduação em Biodiversidade Vegetal
e Meio Ambiente pela infraestrutura e suporte oferecidos.
Ao Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) pela
concessão da bolsa de doutorado.
À Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) pelo apoio
financeiro aos projetos “Basidiomycota (Agaricales e Aphyllophorales) do Parque Estadual
das Fontes do Ipiranga” (processo FAPESP 04/04319-2) e “Agaricales da Reserva Biológica
de Paranapiacaba” (processo FAPESP 09/53272-2).
À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) pelo
apoio financeiro ao projeto “Mata Atlântica e Floresta Amazônica: diversidade e
potencialidades de Agaricales, Polyporales, gasteromicetos e fungos zoospóricos”
(CAPES/PNADB 23038.000043/2010-43).
À minha orientadora, Dra. Marina Capelari, por todo apoio, confiança, disponibilidade
e dedicação para realização deste trabalho. Minha gratidão vai além do que está escrito nessas
folhas pois lhe tenho como um exemplo a ser seguido. Tenha certeza que cada ensinamento
transmitido ao longo desses sete anos será levado por toda minha vida profissional. Muito
obrigado!!!
À Dra. Noemia Kazue Ishikawa, Instituto Nacional de Pesquisas da Amazônia, pela
parceria no projeto CAPES / PNADB, pelo acolhimento em Manaus e por me proporcionar
bons momentos na Floresta Amazônica. Meu muito obrigado também por ter me trazido à
São Paulo para conhecer o trabalho de um taxonomista. Dez anos se passaram e seu incentivo
parece que deu certo!
Ao Dr. Iuri Goulart Baseia, Universidade Federal do Rio Grande do Norte, pela
parceria no projeto CAPES / PNADB e pelo bons momentos durante as coletas em Natal e
região.
Aos membros da banca do exame de qualificação, Dra. Vera Lúcia Ramos Bononi,
Dra. Sandra Farto Botelho Trufem e Dr. Denilson Peralta, pelas sugestões durante o
desenvolvimento deste trabalho.
Aos membros da banca examinadora da defesa de tese, Dr. Vagner Gularte Cortez, Dr.
Aníbal Alves de Carvalho Júnior, Dra. Rosely Ana Piccolo Grandi, Dra. Iracema Helena
Schoenlein-Crusius e Dra. Marina Capelari pelas sugestões para melhoria da versão final
deste trabalho.
A todos os professores do Programa de Pós-graduação em Biodiversidade Vegetal e
Meio Ambiente pelos conhecimentos transmitidos e a excelente formação proporcionada.
A todos os funcionários do Instituto de Botânica, em especial aos da Biblioteca e da
secretaria da Pós-graduação, por toda presteza e auxílio durante a realização deste trabalho.
A todos os pesquisadores, funcionários e amigos do Núcleo de Pesquisa em Micologia
pelos bons momentos e valiosas contribuições.
Aos curadores dos herbários ARAN, BAFC, BPI, BR, F, FH, ICN, INPA, JPB, K,
LIL, MBM, MICH, NY, NYS, PACA, RB, RSPF, SIU, SMDB e SP pelo empréstimo dos
materiais para estudo.
To Dr. Alfredo Justo for his assistance during all steps involving the molecular
systematics of the genus Pluteus. I’m also grateful for his constant availability and for his
warm reception during my stay in Worcester.
To Dr. Andrew Minnis for his assistance in many discussion about the taxonomy of
some species occurring in the USA.
To all people from the Hibbett Lab for the special moments in the Clark University,
Worcester: Alexis Carlson, Alfredo Justo, Dimitrios Floudas, Jaya Sathiya Seelan, Laszlo
Nagy, Mitchell Nuhn, Otto Miettinen, Rachael Martin and Romina Gazis. My special
gratitude to Dr. David S. Hibbett for giving me a great learning experience in his lab.
À Fernanda Karstedt, por todo apoio na coleta de materiais em diversas regiões do
Brasil e pela companhia durante esses anos. Pluteus kartedtianus é apenas uma singela
homenagem diante de toda sua contribuição para a base deste trabalho.
Ao Jadson José Souza de Oliveira, pela companhia no laboratório, por sua
contribuição na coleta de alguns materiais e pelo acolhimento em Natal durante as coletas
pelo PNADB.
À Tatiane Asai por sua amizade e constante disponibilidade nas revisões dos textos e
em discussões aprofundadas de grande valia.
Ao Luiz Antonio Silva Ramos por toda dedicação despendida para criação da
belíssima capa.
Ao Klei Sousa pelo excelente trabalho no preparo das ilustrações.
À minha aluna Flávia Ribeiro Santos pelo auxílio na obtenção de sequências de DNA
durante a execução de seu projeto de Iniciação Científica com apoio financeiro pela concessão
de bolsa do Instituto Federal de Educação, Ciência e Tecnologia de São Paulo (IFSP).
A todos os professores e amigos do IFSP pelo companheirismo e amizade!
Ao amigo Hilário Rocha pelo apoio, companheirismo e pelos ótimos momentos
compartilhados desde o início desta etapa.
Ao grande amigo Anderson Luiz dos Santos, meu amigo do coração, por estar sempre
ao meu lado e disposto a ajudar.
À minha amiga Cristiane de Almeida Nascimento pela auxílio durante as etapas
iniciais deste projeto e amizade ao longo desses anos.
À minha grande amiga Esther do Lago e Pretti, amiga de todas as horas com quem
compartilho momentos valiosos.
À minha valiosa família: minha mãe Alda Cristina Facion Menolli, meu pai Nelson
Menolli, meu irmão Marcelo Menolli e minha irmã Simone Cristina Menolli. Obrigado por
estarem comigo nessa conquista. Amo vocês!!!
A todas as pessoas que, de alguma forma, contribuíram para a realização deste
trabalho, minha eterna gratidão.
“Feliz aquele que transfere o que sabe e aprende
o que ensina”.
Cora Coralina
RESUMO
Pluteus é o gênero mais representativo de Pluteaceae e também o com maior número de
registros para o Brasil, porém, na maioria, os registros são incompletos e duvidosos. Baseado
em dados moleculares e em características da superfície pilear e dos pleurocistídios, o gênero
está dividido em três seções (Pluteus, Celluloderma e Hispidoderma), todas elas com
representantes em território brasileiro. Este estudo foi feito com base em análises
morfológicas de materiais depositados em herbários ou a partir de novas coletas em diversas
regiões e também, sempre que possível, a partir da revisão de espécimes-tipo ou da obtenção
de sequências de DNAr (ITS) para os estudos moleculares. Foram listadas 333 exsicatas de
Pluteus, porém 50 materiais não foram localizados ou estavam impossibilitados para
empréstimo. Outras 115 coleções estavam em condições insuficientes para estudo ou
representam espécies de outros gêneros. Foram reconhecidas 48 espécies, incluindo nove
novos registros e 11 espécies novas, distribuídas em 11 estados brasileiros (Amazonas, Bahia,
Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul,
Rondônia, Santa Catarina e São Paulo). A revisão morfológica das coleções brasileiras de
P. amazonicus, P. aquosus, P. fallax e P. paraensis não foi possível, pois os materiais não
foram localizados ou estavam mal preservados. Mesmo assim, essas espécies foram
consideradas como ocorrentes no Brasil devido à descrição diagnóstica apresentada pelos
autores. A seção Celluloderma é a que tem maior representatividade, com 32 espécies,
incluindo dez espécies novas para ciência (P. anomocystidiatus, P. aureolatus,
P. brunneocrinitus, P. cebolinhae, P. crassus, P. crinitus, P. hispidulussimilis,
P. karstedtianus, P. necopinatus e P. paucicystidiatus) e cinco novos registros para o país
(P. eludens, P. fusconigricans, P. homolae, P. sapiicola e P. striatocystis). A seção
Hispidoderma está representada por nove espécies, incluindo quatro novos registros
(P. argentinensis, P. chusqueae, P. maculosipes e P. velutinus) e uma espécie nova
(P. neochrysaegis), enquanto que a seção Pluteus está representada por sete espécies, sendo
todas elas previamente publicadas para o país e apenas revisadas neste trabalho. A obtenção
de sequências de ITS foi possível para 26 das 48 espécies, sendo que oito das 11 espécies
novas são propostas com suporte molecular, possibilitando a inserção de sequências de
materiais brasileiros na filogenia global do gênero e contribuindo para estabelecer relações
entre espécimes geograficamente separados. Uma chave de identificação para as espécies
também é apresentada.
Palavras-chave: Agaricales, Basidiomycota, Pluteaceae, Taxonomia, ITS
ABSTRACT
Pluteus is the most representative genus in Pluteaceae and also the most recorded from
Brazil, but its knowledge includes many incomplete and dubious records. Based on molecular
data, on the structure of the pileipellis and on characteristics of the pleurocystidia, the genus is
divided into three sections (Pluteus, Celluloderma and Hispidoderma), all of them occurring
in Brazilian territory. This study was conducted with morphological analyses of materials
deposited in Brazilian and foreign herbaria and from new collections from different regions of
Brazil, and also, whenever possible, revising the type specimens and obtaining DNAr (ITS)
sequences for molecular studies. A total of 333 exsiccates of Pluteus were listed, but 50
materials were not located or were unavailable for loan. Other 115 collections were too
insufficient for study or represent species of other genera. Forty eight species were
recognized, including nine that are new records and 11 that are newly proposed, distributed in
11 states of Brazil (Amazonas, Bahia, Minas Gerais, Paraná, Pará, Rio de Janeiro, Rio Grande
do Norte, Rondônia, Santa Catarina e São Paulo). The morphological review of the Brazilian
collections of P. amazonicus, P. aquosus, P. fallax and P. paraensis was not possible because
the materials were not located or were in too bad conditions; even so, these species were
considered as occurring in Brazil due to the complete and diagnostic description presented by
the authors. The section Celluloderma has the higher number of species, with 32 records,
including ten species new to science (P. anomocystidiatus, P. aureolatus, P. brunneocrinitus,
P. cebolinhae, P. crassus, P. crinitus, P. hispidulussimilis, P. karstedtianus, P. necopinatus e
P. paucicystidiatus) and five new records for the country (P. eludens, P. fusconigricans,
P. homolae, P. sapiicola e P. striatocystis); section Hispidoderma is represented by nine
species, including four new records (P. argentinensis, P. chusqueae, P. maculosipes e
P. velutinus) and one new species (P. neochrysaegis); and seven species belong to section
Pluteus, all of them previously published from Brazil, and only revised in this study.
Sequences of ITS were obtained for 26 species, including eight of the 11 new species
proposed. The inclusion of the new generated sequences into the global phylogeny of Pluteus
has contributed to the understanding of the relationships between specimens with
intercontinental distribution. In addition, an identification key for all the species considered in
Pluteus is presented.
Palavras-chave: Agaricales, Basidiomycota, Pluteaceae, Taxonomia, ITS
SUMÁRIO
APRESENTAÇÃO ................................................................................................................ xi
INTRODUÇÃO GERAL ....................................................................................................... 1
REFERÊNCIAS BIBLIOGRÁFICAS ................................................................................. 6
CAPÍTULO I: One hundred and thirteen years of Pluteus knowledge in Brazil: revision of
the first collections studied by P.C. Hennings and J. Rick ..................................................... 11
CAPÍTULO II: Phylogeny of three widely distributed species of Pluteus (Pluteaceae,
Agaricales): taxonomic and biogeographic implications ....................................................... 48
CAPÍTULO III: Contributions to Pluteus (Pluteaceae, Agaricales) in Brazil: 21 taxa from
the state of Paraná including three new species ..................................................................... 77
CAPÍTULO IV: Pluteus sect. Hispidoderma in Brazil with new records based on
morphological and molecular data ....................................................................................... 147
CAPÍTULO V: Pluteus sect. Celluloderma in Brazil: phylogeny and taxonomy including
eight new species .................................................................................................................. 173
DISCUSSÃO GERAL ........................................................................................................ 222
APÊNDICES ....................................................................................................................... 224
APÊNDICE A: Lista comentada das espécies de Pluteus citadas para o Brasil ................ 225
APÊNDICE B: Chave de identificação para as espécies de Pluteus ocorrentes no Brasil . 231
APÊNDICE C: Lista das coleções brasileiras de Pluteus depositadas em herbário ........... 237
CRÉDITO DAS IMAGENS ............................................................................................... 261
N. Menolli Jr., 2013. O gênero Pluteus no Brasil: revisão taxômica e contribuição à filogenia molecular
xi
APRESENTAÇÃO
Este trabalho constitui uma revisão taxonômica das espécies de Pluteus que ocorrem
no Brasil. A partir da revisão morfológica de materiais herborizados e da identificação de
materiais recém coletados em diversas regiões foi possível obter a atualização do
conhecimento do gênero no país. Estudos moleculares baseados em sequências da região ITS
(Internal Trancribed Spacer) do DNAr contribuíram para a filogenia do gênero e o
reconhecimento de parte das espécies aqui apresentadas.
Os resultados deste trabalho estão apresentados em cinco capítulos, redigidos em
inglês, na forma de artigo completo, e formatados de acordo com as normas de publicação das
revistas aos quais foram ou serão submetidos para publicação. Cada capítulo contém uma
seção própria destinada aos materiais e métodos para a realização dos estudos. Os capítulos
apresentados são precedidos de uma introdução geral sobre a família Pluteaceae com especial
atenção à taxonomia e filogenia do gênero Pluteus, bem como seu conhecimento no país. Ao
final é apresentada uma discussão geral dos resultados obtidos e, nos apêndices, uma lista
comentada das espécies de Pluteus citadas para o Brasil (APÊNDICE A), uma chave de
identificação de todas as espécies consideradas certamente conhecidas para o país
(APÊNDICE B) e uma lista das coleções brasileiras de Pluteus depositadas em herbário
(APÊNDICE C).
O capítulo I apresenta a revisão morfológica das primeiras coletas brasileiras de
Pluteus estudadas por P.C. Hennings e J. Rick no início do século passado. Este artigo foi
aceito para publicação na revista Mycotaxon.
O capítulo II apresenta um detalhado estudo morfológico e molecular de três espécies
de Pluteus com distribuição intercontinental. Este artigo foi submetido para publicação na
revista Organisms Diversity & Evolution.
O capítulo III apresenta o estudo morfológico das espécies de Pluteus coletadas no
estado do Paraná entre 1979 e 2011 por André A.R. de Meijer. No artigo constam as
descrições de 21 espécies, incluindo sete novos registros e três novas espécies para o Brasil.
Este artigo será submetido para publicação na revista Nova Hedwigia.
O capítulo IV apresenta uma síntese das espécies de Pluteus seção Hispidoderma
conhecidas para o Brasil. Com base em dados morfológicos e moleculares são apresentados
dois novos registros para o país. Este artigo será submetido para publicação na revista
Cryptogamie Mycologie.
O capítulo V apresenta uma síntese das espécies de Pluteus seção Celluloderma
conhecidas para o Brasil. Com base em dados morfológicos e moleculares são apresentadas
N. Menolli Jr., 2013. O gênero Pluteus no Brasil: revisão taxômica e contribuição à filogenia molecular
xii
oito espécies novas para o país. Este artigo será submetido para publicação na revista
Mycologia.
Os epítetos das espécies novas aqui descritas serão validamente publicados em outro
local, de modo que esta publicação não deve ser considerada para efeitos taxonômicos,
conforme orientado pelos artigos 30.8 e 36.1 do Código Internacional de Nomenclatura
Botânica para algas, fungos e plantas de Melbourne (McNeill et al. 2012). Tais nomes são
disponibilizados aqui para melhor apresentação dos resultados e correta organização do texto.
Quando da submissão dos artigos serão também providenciados os números de registro no
MycoBank para as novas espécies e, se necessário, uma diagnose separada em latim ou inglês.
As sequências de DNA obtidas neste trabalho e mencionadas nos capítulos IV e V
serão depositadas no GenBank quando da submissão dos artigos para publicação. Sendo
assim, as novas sequências de DNA apresentadas nesses capítulos estão, até o momento,
identificadas apenas pelo número de coletor ou número de acesso de herbário.
Para melhor organização do texto, os capítulos estão organizados na ordem em que
espera-se que sejam publicados, possibilitando que os dados de um capítulo precedente sejam
citados em um capítulo ulterior. Nesses casos a citação ocorre na forma de parte/capítulo de
uma tese conforme seção 7.3 da NBR 6023 (ABNT 2002).
N. Menolli Jr., 2013. O gênero Pluteus no Brasil: revisão taxonômica (...)
Introdução Geral - 1
INTRODUÇÃO GERAL
O gênero Pluteus Fr., tipificado por P. cervinus (Schaeff.) P. Kumm, contém
aproximadamente 300 espécies de ampla distribuição mundial e é o mais representativo da
família Pluteaceae Kotl. & Pouzar (Kirk et al. 2008). De acordo com a classificação adotada
por Pegler (1983) e Singer (1986), Pluteaceae é constituída de três gêneros: Chamaeota
(W.G. Smith) Earle, Pluteus e Volvariella Speg. que juntos totalizam 359 espécies (Kirk et al.
2008) e compartilham as seguintes características ecológicas e morfológicas: hábito terrestre,
humícola, lignícola, coprófilo ou raramente micoparasita, porém, nunca ectomicorrízico;
basidiomas carnosos ou membranosos, com ampla variação de tamanho; lamelas livres e
rosadas; esporada rosada; estipe central e bem desenvolvido; volva e anel presentes ou
ausentes; trama do himênio bilateral convergente; basidiósporos globosos a elípticos, de
parede espessada, lisos, inamiloides e cianófilos; cistídios geralmente presentes, metuloides
ou não (Pegler 1983, Orton 1986, Singer 1986). Kirk et al. (2008) consideraram quatro
gêneros no verbete Pluteaceae, porém, o quarto gênero não foi encontrado na publicação.
Tradicionalmente, as esp cies de Pluteus s o distingu veis dos demais g neros de
Pluteaceae, basicamente, pela aus ncia de volva e anel, uma vez que Chamaeota é
caracterizado, dentre outros fatores, pela presença de anel e ausência de volva, e Volvariella
pela presença de volva e ausência de anel (Pegler 1983, Orton 1986, Singer 1986). Entretanto,
essa delimita o vem sendo uestionada nos ltimos anos com base em estudos morfológicos
e moleculares (Minnis et al. 2006, Corriol & Moreau 2007, Malysheva et al. 2009, Justo et al.
2011a, b, Vizzini & Ercole 2011).
Minins et al. (2006), baseados em dados moleculares (nLSU), propuseram a
combinação de Chamaeota mammilata (Longyear) Murrill em Pluteus mammillatus
(Longyear) Minnis, Sundb. & Methven, aceitando, portanto, a inclusão de espécies com anel
no gênero Pluteus e tornando o reconhecimento de Chamaeota como artificial, já que, de
acordo com as análises moleculares, essa espécie está inserida em um clado junto com outras
espécies de Pluteus. A relação entre Chamaeota e Pluteus, bem como a delimitação de
Chamaeota com base na presença de anel, já foi discutida anteriormente por Singer (1986),
que ressaltou a impossibilidade, até aquele momento, de confirmar a distinção dos dois
gêneros apenas pela ausência de anel. Posteriormente, a partir de dados morfológicos e/ou
moleculares, outros autores passaram a aceitar no gênero Pluteus espécies de Pluteaceae com
anel (Corriol & Moreau 2007, Malysheva et al. 2009, Justo et al. 2011a, b, Vizzini & Ercole
2011).
O posicionamento de Volvariella na família Pluteaceae também tem sido alvo de
estudos moleculares recentes. A partir de análises filogenéticas utilizando sequências do gene
nLSU, Moncalvo et al. (2002) demonstraram o posicionamento de espécies de Volvariella
[V. volvacea (Bull.) Singer e V. hypopithys (Fr.) Shaffer] distante do clado pluteoide e
agrupado, embora sem suporte estatístico, à sequências de Fistulina Bull. e Schizophyllum Fr.
Posteriormente, a partir de uma abordagem multigênica com representantes de Agaricales,
Matheny et al. (2006) demonstraram a posição de V. gloiochepala (DC.) Boekhout & Enderle
como grupo irmão de Pluteus em um clado com suporte estatístico. Com base nesses
resultados, Justo et al. (2011a) conduziram um estudo filogenético de Pluteaceae e
propuseram um novo g nero (Volvopluteus Vizzini, Contu & Justo) para as espécies
posicionadas como grupo irmão de Pluteus, uma vez que Volvariella sensu lato foi
considerado polifilético. Além dos dados moleculares, de acordo com Justo et al. (2011a),
Volvariella e Volvopluteus podem ser distinguidos pelo comprimento dos basdiósporos (maior
que 11 µm em Volvopluteus) e pela estrutura da superfície pilear (indiferenciada, gelatinizada
e composta por hifas menores que 15 µm de diâmetro em Volvopluteus e não-gelatinizada,
indiferenciada ou quase tricodérmica e composta por hifas maiores que 15 µm de diâmetro em
Volvariella).
Atualmente, portanto, a delimitação molecular de Pluteaceae (clado Pluteoide) inclui
como monofiléticos os gêneros Pluteus (incluindo Chamaeota) e Volvopluteus, e como grupo
irmão Melanoleuca Pat. (Tricholomataceae R. Heim ex Pouzar). Volvariella, por sua vez, não
tem a mesma origem filogenética que Pluteus e Volvopluteus e parece estar relacionada com
Cantharocybe H.E. Bigelow & A.H. Sm. e Camarophyllus (Fr.) P. Kumm. (Justo et al.
2011a).
Chamaeota e Volvariella s o grupos de conhecimento e registros mais escassos para o
Brasil quando comparados à Pluteus sensu Singer (1986). Chamaeota compreende
apro imadamente dez esp cies (Minnis et al. 200 , por m sem nenhum registro para o
Brasil, enquanto que Volvariella sensu lato compreende apro imadamente 50 esp cies (Kirk
et al. 2008), sendo 19 registradas para o Brasil (Menolli & Capelari 2008), incluindo duas
espécies agora combinadas em Volvopluteus. Por outro lado, Pluteus, sendo o g nero mais
bem representado de Pluteaceae, é também o de maior número de registros para o Brasil,
incluindo de 45 a 70 nomes já mencionados para o país, de acordo os levantamentos de
Putzke & Wartchow (2008) e Menolli et al. (2010), respectivamente.
As espécies de Pluteus possuem hábito preponderantemente lignícola, crescendo
frequentemente sobre madeira em decomposição, podendo ser, às vezes, humícolas ou
terrestres (Pegler 1983, Orton 1986, Singer 1986). Uma vez que, atualmente, as espécies de
Pluteaceae com anel estejam também incluídas no gênero Pluteus, a ausência de volva passa
a ser a principal característica morfológica para o reconhecimento de Pluteus dentro da
família. A presença de cistídios metuloides frequentemente com prolongamentos apicais e a
organização da superfície pilear constituída por elementos globosos ou por hifas alongadas
são também características úteis para a organização infragenérica e o reconhecimento das
espécies de Pluteus (Singer 1986).
A classificação infragenérica mais aceita para Pluteus foi proposta por Singer (1959,
1986) e se baseia na divisão do gênero em três seções (Pluteus, Hispidoderma Fayod e
Celluloderma Fayod) com base em características morfológicas, como a estrutura da
superfície pilear e características dos pleurocistídios. Dessa forma, uma camada himeniforme
de células globosas ou subglobosas caracteriza a superfície pilear da seção Celluloderma,
enquanto que as outras duas seções exibem a superfície pilear com hifas alongadas, dispostas
de forma ereta, subereta ou prostradas e diferem entre si pela presença de pleurocistídios
metuloides na seção Pluteus ou de parede fina na seção Hispidoderma (Singer 1959, 1986).
A seção Celluloderma, por sua vez, é dividida em duas subseções: a subseção
Eucellulodermini Singer, que apresenta a superfície pilear com elementos uniformes e
dermatocistídios ausentes, e a subseção Mixtini Singer que possui dermatocistídios e
elementos dimórficos constituindo a superfície pilear (Singer 1956, 1959, 1986).
Vellinga & Schreurs (1985) propuseram uma variação dessa classificação, porém sem
grande aceitação. Nessa classificação, os autores também mantém a divisão do gênero em três
seções com base na estrutura da superfície pilear e nas características dos pleurocistídios.
Duas das três seções possuem espécies com superfície pilear indiferenciada, constituída por
hifas prostradas ou com os elementos terminais ascendentes: a seção Pluteus, mantida de
acordo com a classificação de Singer (1959, 1986), que inclui espécies com pleurocistídios
metuloides, e a seção Villosi Schreurs & Vellinga, proposta para acomodar parte dos táxons
anteriormente classificados na seção Hispidoderma (Singer 1959, 1986) e composta por
espécies com pleurocistídios ausentes ou, se presentes, de parede fina e sem prolongamentos
apicais. A seção Celluloderma foi mantida, porém com a inclusão de uma nova subseção
[Hispidodermini (Fayod) Vellinga & Schreurs] para acomodar o restante das espécies da
seção Hispidoderma sensu Singer (1959, 1986). Dessa maneira, seção Celluloderma, de
acordo com Vellinga & Schreurs (1985), engloba uma gama maior de variação da superfície
pilear quando comparada à classificação de Singer (1959, 1986) e inclui espécies que
possuem pleurocistídios de parede fina, superfície pilear tricodérmica, himeniforme ou
intermediária entre himeniforme e epitelial, organizadas em três subseções: Eucellulodermini,
que inclui espécies com a superfície pilear formada por elementos uniformes,
esferopedunculados a clavados; Mixtini, para espécies com superfície pilear constituída de
elementos dimórficos com dermatocistídios; e Hispidodermini, para espécies com superfície
pilear tricodérmica ou himeniforme composta por elementos uniformes, cilíndricos a
fusiformes.
Estudos moleculares recentes (Menolli et al., 2010; Justo et al., 2011a, b) têm
mostrado suporte para a classificação de Singer (1959, 1986), porém com alguns rearranjos.
Justo et al. (2011a) mostraram, a partir de análises filogenéticas da região ITS, que as
espécies com pleurocistídios não metuloides e caracterizadas por uma superfície pilear
indiferenciada composta por hifas prostradas (= Villosi ou Hispidoderma pro parte), aneladas
(= Chamaeota) ou não, estão posicionadas com as demais espécies da seção Celluloderma ao
invés de com aquelas da seção Hispidoderma, ou em uma seção separada (Villosi) como
proposto por Vellinga & Schreurs (1985). Na seção Celluloderma foram também incluídas as
espécies com pleurocistídios não metuloides e superfície pilear epitelial ou himeniforme de
elementos curtos, entremeados ou não (subseção Eucellulodermini + subseção Mixtini) por
elementos cistidioides alongados; entretanto, sem nenhum suporte estatístico para qualquer
subdivisão da seção de acordo com os diferentes tipos de superfície pilear (Justo et al. 2011a).
A seção Hispidoderma, por sua vez, foi caracterizada por incluir espécies com superfície
pilear tricodérmica ou himenidérmica de elementos alongados e bastante variáveis em forma e
tamanho, enquanto que a seção Pluteus ficou restrita às espécies com pleurocistídios
metuloides (ou raramente de parede fina a levemente espessada) e superfície pilear
indiferenciada composta por hifas prostradas (Justo et al. 2011a).
Os trabalhos de Justo et al. (2011a, b) foram, sem dúvida, os mais importantes para a
elucidação da filogenia do gênero. Outros estudos moleculares com espécies de Pluteus estão
centrados no posicionamento de novas espécies ou novos registros (Malysheva et al. 2009,
Rodríguez et al. 2009, 2010; Vizzini & Ercole 2011; Justo et al. 2011c, 2012, Pradeep et al.
2012). Para o Brasil e toda a região sul-americana, os estudos moleculares estão restritos ao
trabalho de Menolli et al. (2010) que apresentou a descrição de dez espécies, incluindo dois
novos táxons e três novos registros para o país. Além disso, Justo et al. (2011a, b) incluíram
dados moleculares de outras oito espécies de Pluteus provenientes do Brasil para compor a
filogenia geral do gênero.
Outros trabalhos que contribuíram para o conhecimento das espécies de Pluteus no
Brasil são baseados em dados morfológicos, incluindo o registro ou descrição de várias
espécies dúbias (Hennings 1900, 1904a, b, Rick 1907, 1919, 1930, 1938, 1961), a
apresentação de espécies em listas ou apenas com uma breve descrição (Bresadola 1920,
Grandi et al. 1984, Singer 1984, Singer & Aguiar 1986, Raithelhuber 1991, Stijve & de
Meijer 1993, Stijve 1995, Pegler 1997, Meijer 2001, 2006, 2009, 2010, Rosa et al. 2003,
Xavier-Santos et al. 2004, Sobestiansky 2005, Drechsler-Santos et al. 2007, Rosa & Capelari
2009, Gugliotta et al. 2012), ou ainda registros e novas espécies apresentadas com descrições
completas dos materiais estudados (Singer 1954, 1956, 1959, 1973, 1989, Singer & Digilio
1952, Wartchow et al. 2004, 2006, Menolli et al. 2010, Menolli & Capelari 2010).
A partir de um cuidadoso levantamento bibliográfico foi possível verificar que o
número de táxons de Pluteus mencionados para o Brasil chega a 85 distribuídos em dez
estados (Amazonas, Bahia, Mato Grosso, Minas Gerais, Par , Paran ,
rande do Sul,
io de aneiro,
io
ondônia e S o Paulo), incluindo 82 nomes e outras cinco espécies não
identificadas (APÊNDICE A). Al m disso, acredita-se ue a am novas esp cies a serem
descritas para o rasil e ue muitos erb rios nacionais e estrangeiros conten am esp cies de
Pluteus ainda n o registradas por meio de publica es cient ficas, o ue contribui para um
n mero incerto do con ecimento do g nero no pa s. Considerando todos os registros
constantes no APÊNDICE A, é possível verificar que o conhecimento do gênero no Brasil é
bastante confuso e necessita de revis o, pois, provavelmente, grande parte dessas cole es
representam identifica es errôneas, sinônimos, esp cies de outros g neros, esp cimes n o
preservados ou que foram apresentados em listas e sem descrição completa para certificação
de sua identificação.
Diante do estado atual de conhecimento das esp cies de Pluteus no Brasil, fez-se
necess rio um estudo de revis o dos materiais
coletados e mencionados em literatura, al m
da realiza o de novas coletas ue pudessem proporcionar a descoberta de esp cies novas e
novas ocorr ncias.
A partir do levantamento das esp cies de Pluteus
registradas para o pa s, bem como
da realização de novas coletas, o propósito deste trabalho foi contribuir para o conhecimento
das espécies de Pluteus a fim de esclarecer problemas ta onômicos e corrigir poss veis erros
de identifica o, bem como estabelecer o posicionamento das coleções brasileiras dentro dos
grupos estudados e contribuir para a filogenia do gênero. O estudo foi feito com base na
análise morfológica de materiais depositados em herbários ou a partir de novas coletas em
diversas regiões e também, sempre que possível, a partir da revisão de espécimes-tipo ou da
obtenção de sequências de DNAr (ITS) para os estudos moleculares.
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Capítulo I - 11
CAPÍTULO I
One hundred and thirteen years of Pluteus knowledge in Brazil: revision of the first
collections studied by P. C. Hennings and J. Rick
Nelson Menolli Jr. & Marina Capelari
Artigo aceito para publicação na revista Mycotaxon
One hundred and thirteen years of Pluteus knowledge in
Brazil: revision of the first collections studied by P. C.
Hennings and J. Rick
NELSON MENOLLI JR.1,2* & MARINA CAPELARI2
Instituto Federal de Educação, Ciência e Tecnologia de São Paulo, Campus São Paulo,
CCT / Biologia, Rua Pedro Vicente 625, 01109-010 São Paulo, SP, Brazil
2
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902
São Paulo, SP, Brazil
* CORRESPONDENCE TO: [email protected]
1
ABSTRACT — The first Brazilian collections of Pluteus studied by P. C.
Hennings and J. Rick during the early 20th century were re-examined,
complementing Brazilian knowledge of the genus. Searches of the
bibliographical and herbarium records led to a total of 32 Pluteus names that
were linked to specimens collected in Brazil and studied by Hennings and Rick.
Ten are represented by Brazilian types, but five of these could not be located in
any of the consulted herbaria, representing, therefore, nomina dubia. Moreover,
previously published collections listed under five other names could not be
located either. All other collections were in BPI, FH, PACA and SP and they
were studied for morphology. In all cases, European names originally attributed
by Hennings and Rick were found to be inappropriate and identifications based
especially on species described from the Neotropics are suggested. Sections
Pluteus and Celluloderma are represented by most collections.
KEY WORDS — biodiversity, herbarium, Pluteaceae, taxonomy
Introduction
Pluteus Fr. is the most representative genus of Pluteaceae (Agaricales,
Basidiomycota), with approximately 300 species worldwide (Kirk et al.
2008). Although Pluteus species are easily recognizable by their free
lamellae, pinkish spore print, inamyloid basidiospores, inverse hymenophoral
trama and the absence of annulus and volva (Singer 1986), modern molecular
studies have contributed to resolving some taxonomic inconsistencies,
establishing synonymies of species described multiple times, and enlarging
the circumscription of the genus to include annulate species historically
classified in the allied genus Chamaeota (W.G. Sm.) Earle (Minnis et al.
2006, Justo et al. 2011ab, Vizzini & Ercole 2011). However, in terms of old
and especially poorly preserved collections, molecular tools are limited in
their ability to resolve taxonomic problems. An accurate morphological study
is an alternative in these situations, although it is not always possible to
retrieve useful information because the hymenial structures in old collections
are frequently collapsed and not revivable.
Capítulo I - 12
Paul Christoph Hennings was the first author to describe a Pluteus species
from Brazil based on material collected by R. Pilger in 1899, which was
thereafter described as P. scruposus Henn. (Hennings 1900). Hennings
(1904ab) also described P. cervinus var. griseoviridis and P. termitum Henn.
based on materials collected by A. Puttemans in 1903 and E.H.G. Ule in
1901, respectively.
Hennings was a German mycologist who studied the fungi from Brazil in
the late 19th and early 20th centuries. His works were based on materials
collected in different regions of Brazil by many collaborators, including E.H.
Ule, F.A.G.J. Möller, A.F.M. Glaziou, C.A.W. Schwacke, I.D.J. Huber, C.F.
Baker, E.A. Goeldi, A. Puttemans, and R. Pilger (Fidalgo 1968). Hennings
was born in 1841 in Heide (Germany) and, from approximately 1885 onward,
he devoted himself to the study of fungi from around the world, mainly those
from German colonies and Brazil. He published more than 250 papers and
brought together in Berlin one of the best collections of fungi in the world
(Perkins 1909). Many of the collections described by Hennings were
deposited in herbarium B of the Botanical Museum Berlin-Dahlem, which
was partly destroyed by fire in a bombing raid in 1943 (Merrill 1943, Hiepko
1987). However, several duplicates of Hennings’ types from different groups
of Fungi (especially Fungi Imperfecti and Uredinales) were saved because
they were also deposited in other herbaria (Hiepko 1987, Hein 1988). Hein
(1988) published a list of all species described by Hennings and data about
the preservation states of these collections in B, but, unfortunately, no
Pluteus collections survived the fire in B.
Among Henning’s collaborators was Ernst Heinrich Georg Ule, an
important and active collector of fungi in Brazil. Ule was born in 1854 in
Halle an der Saale (Germany), and in 1883 he moved to Brazil, first living in
Santa Catarina State and then in Rio de Janeiro State, where he became a
traveling naturalist of the National Museum (Martius 1906, Fidalgo 1968).
The specimen of P. termitum described by Hennings (1904a) was collected
during the expedition of Ule (19.X.1901 – 18.XI.1901) to the Amazonian
Forest in Fortaleza, Amazonas State (Hennings 1904a, Martius 1906). Ule
accumulated approximately 3,361 species of fungi from the Brazilian States
of Santa Catarina, Rio de Janeiro, Minas Gerais, Goiás and Amazonas
(Martius 1906). These collections were distributed to many mycologists,
including G. Winter, F.O. Pazschke, H. Rehm, H. Sydow, P. Sydow, E. Jahn,
P. Dietel, and P. C. Hennings (Fidalgo 1968).
Other collaborators in Henning’s Pluteus studies were Robert Pilger and
Arsène Puttemans. Pilger was born in 1876 in the Helgoland Archipelago
(Germany), and in 1899 he conducted his expeditions to Mato Grosso State,
where he collected the first Pluteus from Brazil. Puttemans was born in
Capítulo I - 13
Brussels (Belgium) in 1873 and he moved to Brazil in 1892, where he lived
first in Rio de Janeiro State for two years, and then in São Paulo State.
Puttemans is undoubtedly the most important name in Brazilian
phytopathology, and he was also an important collector of various groups of
non-pathogenic fungi. Although many other collections sent to Germany
were destroyed in the fire in B, fortunately, duplicates of almost all of
Puttemans’ collections were deposited in other herbaria, and approximately
7,000 samples were preserved at UFRJ of the ‘Universidade Federal Rural do
Rio de Janeiro’ (Fidalgo 1968); additional collections of Puttemans are also
preserved in other herbaria, including the type of P. cervinus var.
griseoviridis that is deposited in herbarium SP of the ‘Instituto de Botânica’.
As the study of Brazilian fungi began with foreign collectors, Johann
Rick played an extremely important role in the history of Brazilian
mycology, and he is considered the father of Brazilian mycology. In 1904, he
started a series of publication of mycological papers representing the
beginning of the systematic study of fungi from Brazil (Fidalgo 1962, 1968).
His collections brought together approximately 12,000-15,000 exsiccatae
from Rio Grande do Sul State that are now deposited in the herbarium PACA
of the ‘Instituto Anchietano de Pesquisas’ (Fidalgo 1968). Rick described and
recorded 21 Pluteus taxa from Rio Grande do Sul State (Rick 1907, 1919,
1930, 1938, 1961), including seven new taxa from Brazil (P. cristatulus Rick,
P. exiguus var. venosus, P. fibrillosus Rick, P. leptonia, P. sensitivus Rick,
P. straminellus, and P. velatus).
Rick was born in Dornbirn (Austria) in 1869, and in approximately 1896
he began to study fungi, and he came into contact with H. Rehm,
G. Bresadola and C.G. Lloyd. He shared many duplicates with Lloyd, which
formed what was most likely the largest collection of Brazilian fungi at the
time, the ‘Lloyd Herbarium’ (Fidalgo 1962). Presently, the collections of the
‘Lloyd Herbarium’ are deposited in the U.S. National Fungus Collections
(BPI), which also includes recent collections and the contributions of many
collectors, giving BPI a total of approximately 14,000 collections of different
groups of fungi from Brazil (Farr & Rossman 2013). Rick arrived in Brazil in
1903 and he was appointed as a professor of the Jesuit College of São
Leopoldo, Rio Grande do Sul State, where he collected most of his specimens
(Fidalgo 1962), including what likely represents his first Pluteus specimen,
which is dated 1906 and was later identified as P. cervinus (Rick 1938,
1961). Rick continued studying Brazilian fungi until 1946, the year he died in
Brazil.
Considering that foreign mycologists made the first studies of Pluteus
from Brazil, many names proposed for European specimens were largely
used for Brazilian materials. Nowadays, the state of knowledge of Pluteus in
Capítulo I - 14
South America is somewhat problematic mainly due to the abundance of
species that have been described for this region (Singer 1959, Horak 1964)
and to inappropriate identifications of the past. Nevertheless, until now, there
is no published paper correcting past identifications. Thus, with the aim of
complementing the study of Pluteus from Brazil, we present a re-examination
of the historical Pluteus collections studied by Hennings and Rick, including
comments about the identifications. Undoubtedly, studies involving a
combination of multiple methods and different types of evidence are relevant
for the knowledge about the evolution of the genus and to determine if
morphologically similar taxa from different continents are conspecific or
distinct. However, in cases where we have only old and poorly preserved
collections, historical studies are equally important to disseminate the
corrections about past mistakes involving European names misapplied to
South American species and to reduce the dissemination of errors, for
instance about the geographical distribution of selected species.
Materials & methods
The materials examined herein include types and other collections of Pluteus
studied by Hennings and Rick that are referenced in their publications (Hennings
1900, 1904ab; Rick 1907, 1919, 1930, 1938, 1961). Unpublished collections that they
studied were also re-examined. The main herbaria consulted to locate as many
collections studied by them as possible were B, BPI, FH, K, KIEL, L, PACA, S, SP
and W. The herbarium acronyms follow Thiers (2013). When necessary, additional
collections were also studied to infer data about the identification of specimens.
The names are arranged alphabetically according to specific epithet attributed by
Hennings and Rick and their protologues are presented. The collections studied by
t em under t e name presented in t e ta onomic eader are listed under ‘specimen
e amined’, and t ose previously publis ed are followed by an asterisk ‘*’. In t e
section ‘additional specimen(s e amined’ are listed t e collections preserved in
herbaria under other names but that share the same characters that the materials listed
in ‘specimen e amined’. Specimens t at do not represent t e identification suggested
for t e ot er materials involved under t is name are also listed in ‘additional
specimen(s e amined’, but in t is case t e correct identification and name are
discussed in the text. For each taxon, comments are presented about the name
attributed by Hennings or Rick and about what these collections actually represent
based on our morphological examination. To summarize the results, the data were
compiled in Table 1. T e section ‘Ta onomy’ is reserved to include data about t e
collections studied, whereas the data for collections that were previously published
but for which the specimens were not located are only listed in Table 1. Generic and
infrageneric concepts follow Singer (1959, 1986) as complemented by Justo et al.
(2011ab).
For microscopic analyses, the dried material was wetted with 70% ethanol and
then rehydrated in 5% KOH or stained wit Melzer’s reagent to determine t e
Capítulo I - 15
Capítulo I - 16
amiloidity reaction of the spores. The notation [a/b/c] at the beginning of a set of
basidiospore data is to be read as “(a basidiospores were measured from (b
basidiomata taken from (c) collections”. Q represents t e range of t e lengt /widt
ratio for all of the measured spores, Qm represents the average of all calculated Q
values for all of the measured basidiospores and Lm (Wm) represents the average of
all of the lengths (widths) of the measured basidiospores. At least 20 basidiospores
from each basidioma were measured in lateral view, and the terms denoting
basidiospore shape follow Bas (1969). For the pileipellis having a hymeniderm, the
length of the pedicel was included in the measurement of the pileipellis elements, and
its length is also reported separately.
Results
A total of 32 names of taxa associated with collections were recovered;
Hennings studied three of them and the other 29 were studied by Rick (Table
1). In addition to the 21 taxa published by Rick (1907, 1919, 1930, 1938,
1961), eight additional names were recovered from his unpublished
materials. Ten taxa are represented by Brazilian types, but five of these were
not located in any herbarium consulted, and we now regard these as nomina
dubia. Moreover, we were unable to locate collections bearing five other
names listed here. Thus, this study comprises a taxonomic revision with
comments about the correct identification of material identified under 22
names. The identifications suggested are based especially on species
described from the Neotropics making European names originally attributed
by Hennings and Rick inappropriate in all cases.
Sections Pluteus and Celluloderma are represented by most collections.
Probably these sections are the most representative in the genus as showed by
the number of sampled collections by Justo et al. (2012b). A current study
(Menolli, de Meijer & Capelari, unpublished data) including the
morphological revision of Pluteus specimens collected during more than 30
years of random field trips in South Brazil also revealed a preponderance of
species in these sections. The preponderance of collections of sections
Celluloderma and Pluteus was also verified in the studies of Singer (1959)
and Horak (1964) in the Neotropics.
Table 1: Pluteus species sensu Hennings and Rick reported from Brazil.
Species (sensu Henn. and Rick)
“P. brunneopictus Berk.”*
References
Rick (1938, 1961)
“P. cervinus Schaeff.”
P. cervinus var. griseoviridisB
Rick (1938, 1961)
Hennings (1904b)
P. cervinus var. patricius (Schulzer) Massee
P. cristatulusB
“P. dissimilis”, nom. in herb.
“P. egregius”, nom. in herb.
“P. exiguus Pat.”
P. exiguus var. venosus, nom. inv.B
Rick (1907)
Rick (1938, 1961)
unpublished
unpublished
Rick (1938, 1961)
Rick (1961)
Comments
most likely P. tucumanus
Singer and P. cubensis
(Murrill) Dennis
P. xylophilus (Speg.) Singer
most likely P. angustisporus
Singer
not found
not found (nomen dubium)
P. xylophilus
not found
most likely P. jamaicensis
Murrill
Table 1, concluded.
Species (sensu Henn. and Rick)
“P. eximius Saund et Sm.”
P. fibrillosusB
“P. fuscidulus”, nom. in herb.
“P. granulatus Bres.”
References
Rick (1919, 1938, 1961)
Rick (1938, 1961
unpublished
Rick (1930, 1938, 1961)
“P. hispidulus Fr.”
Rick (1919, 1938, 1961)
“P. leoninus Schaeff”
Rick (1938, 1961)
B
P. leptonia
“P. longistriatus Peck”
Rick (1938, 1961)
unpublished
“P. melanodon Secr.”
“P. murinus Bres.”
Rick (1938, 1961)
unpublished
“P. nanus Pers.”
Rick (1919, 1938, 1961)
P. nanus var. podospileus (Sacc. & Cub.)
Rick
“P. pellitus Pers.”*
Rick (1938, 1961)
Capítulo I - 17
Comments
not found
too insufficient to study
most likely P. glaucotinctus
E. Horak
most likely P. yungensis
Singer
most likely P. conizatus
(Berk. & Broome) Sacc.
Entoloma s.l.
most likely a true P.
longistriatus
not found
P. albostipitatus (Dennis)
Singer
most likely P. sapiicola
Singer
not found
Rick (1930, 1938, 1961)
most likely P. petasatus (Fr.)
Gillet or a species in sect.
Hispidoderma
“P. princeps”, nom. in herb.
unpublished

“P. phlebophorus Dittm.”*
Rick (1938 , 1961)
most likely P. sapiicola and
P. tucumanus
B

P. scruposus Henn.
Hennings (1900 )
B



P. sensitivus
Rick (1930 , 1938 , 1961 )
“P. sensitivus var. macrospora Rick”, nom.
unpublished
P. albostipitatus (Dennis)
in herb.
Singer
B
P. straminellus
Rick (1961)
nomen dubium
B

P. termitum
Hennings (1904a )
“P. umbrosus Pers.”
Rick (1919, 1938, 1961)
B
P. velatus
Rick (1961)
nomen dubium
“P. wehlianus Müller”
unpublished
Bolbitiaceae or
Strophariaceae
B
= type locality in Brazil; * = species represented by two collections that are actually different species (see
discussion in the text);  = reference without indication of the collection studied and/or the herbarium where
the materials are deposited.
Taxonomy
“Pluteus brunneopictus Berk.” sensu Rick, Iheringia Bot. 8: 420. 1961.
FIGURE 1
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Santa Maria city,
1936, J. Rick s.n. (PACA14522*).
MICROCHARACTERS — BASIDIOSPORES [20/1/1] 5.0–6.2(–7.5) × 3.7–5.0 µm
[Q = 1.24–1.35(–1.50); Qm = 1.31; Lm = 5.7 µm; Wm = 4.4 µm], broadly
ellipsoid to ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate.
PILEIPELLIS a hymeniderm, composed of spheropedunculate or clavate to
clavate-pedicellate cells, 27–40(–50) × 11.2–18.7(–22) µm, with a medium to
long pedicel (5.0–17.5 µm long), thin-walled, with brown internal content,
frequently condensed or sometimes dissolved. P LEURO- AND CHEILOCYSTIDIA
not recovered. CLAMP CONNECTIONS absent.
ADDITIONAL SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, São
Leopoldo city, 1943, J. Rick s.n. (PACA14531).
COMMENTS — Pluteus brunneopictus (Berk. & Broome) Sacc. is an obscure
species described from Sri Lanka based on one sample collected in 1868
(Berkeley & Broome 1871, Saccardo 1887). Although there are no recent
collections, Singer (1956) and Pegler (1986) re-examined the type and
confirmed the occurrence of broadly ellipsoid to ellipsoid basidiospores (5.3–
7.3 × 4.5–4.8 µm measured by Singer and 7.0–9.0 × 5.0–6.0 µm measured by
Pegler) and a cellular pileipellis composed of ellipso-pedicellate elements
with brownish pigment. These data are similar to what we observed in the
material studied by Rick (1938, 1961), but they are not enough (mainly
because pleuro- and cheilocystidia were not found) to correlate it to
P. brunneopictus.
FIGURE 1. Pluteus brunneopictus sensu Rick (PACA14522). a. Basidiospores; b. Pileipellis
cells. Scale bars = 10 µm.
Moreover, considering the geographical distance, it is unlikely that the
specimen collected by Rick in South America is actually P. brunneopictus.
Similarly, Singer (1959) suggested that P. brunneopictus sensu Rick might be
P. umbrinoalbidus Singer, P. rimosoaffinis Singer, or any other species of
stirps Pulverulentus, Jamaicensis or Tucumanus. Based on the insufficient
information available, because the hymenial structures were not recovered,
we consider that this collection is most likely P. tucumanus, originally
described from Argentina. According to Singer (1959), P. tucumanus has
ellipsoidal basidiospores (3.8–5.5 × 2.7–3.8 µm) and subglobose-pedicellate
pileipellis cells with dissolved or condensed intracellular pigment, and it
Capítulo I - 18
Capítulo I - 19
represents one of the few South American species in sect. Celluloderma that
is characterized by a predominantly brown color in the pileus and by ellipsoid
basidiospores. More collections of Pluteus from Brazil and Argentina with
these characteristics would be required to establish the actual relationship
between ick’s collection and Singer’s species, and to confirm its occurrence
in Brazil.
It is also important to record that Rick (1961) mentioned under
P. brunneopictus two collections (PACA14522 and PACA14532) made in
Santa Maria city in 1936. However, the exsiccata deposited at PACA and
mistakenly labeled (PACA)14532 actually contains the collection
PACA14531 (São Salvador city, 1943). PACA14531 (Fig. 2) represents an
unpublished sample and, according to our re-examination, it has broadly
ellipsoid to rarely ellipsoid basidiospores, [20/1/1] 6.2–7.5(–8.7) × 5.0–6.2
(–7.5) µm [Q = (1.16–)1.21–1.24(–1.50); Qm = 1.24; Lm = 7.1 µm; Wm =
5.7 µm]; clavate or elongate-ventricose pleurocystidia (47–68 × 15.0–22
µm); pileipellis that seems to be a cutis with ascendant elements, and clamp
connections are absent. Cheilocystidia were not found. Based on these
characteristics and on a current study (Menolli, de Meijer & Capelari,
unpublished data) that includes a broad examination of collections of
P. cubensis, this seems to be the better name for PACA14531. Pluteus
cubensis is a widely distributed species in Central and South America, and its
representatives display wide morphological variation, mainly regarding the
shape and size of the basidiospores.
FIGURE 2. Pluteus brunneopictus
b. Pleurocystidia. Scale bars = 10 µm.
sensu
Rick
(PACA14531).
a.
Basidiospores;
“Pluteus cervinus Schaeff.” sensu Rick, Iheringia Bot. 8: 418. 1961. FIGURE 3
SPECIMENS EXAMINED — Brazil, Rio Grande do Sul State, São Leopoldo city,
1906, J. Rick s.n. (PACA14519*); 1934, J. Rick s.n. (PACA14516*); 1934,
leg. Steffen, det. J. Rick (PACA14526).
MICROCHARACTERS — BASIDIOSPORES [62/4/4] (5.0–)6.2–7.5(–8.7) × (3.7–)
5.0–6.2 µm [Q = (1.21–)1.24–1.68; Qm = 1.45; Lm = 6.7 µm; Wm = 4.8
µm], broadly ellipsoid, ellipsoid to elongate, inamyloid, hyaline, smooth,
thick-walled, guttulate. P LEUROCYSTIDIA of three types: (I) the normal
Cervinus-type, 58–76 × 15.0–17.5 µm, fusoid-ventricose, colorless, hyaline,
with four to six apical to lateral prongs at the apices, thick-walled, the wall
sometimes tapering toward the base, abundant to very abundant; (II) the
modified Cervinus-type as the majority of those found in P. harrisii Murrill,
56–87 × 12.5–20 µm, fusoid-ventricose, colorless, hyaline, usually without
prongs and a subcapitate apex or with two short lateral prongs and slightly
strangulated at the apex, moderately thick-walled, common; and (III) the
Magnus-type, 56–78 × 12.5–21 µm, lageniform to fusoid, with an acute apex,
colorless, hyaline, moderately thick-walled, rare. CHEILOCYSTIDIA not
recovered. PILEIPELLIS a cutis. CLAMP CONNECTIONS absent.
ADDITIONAL SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Pareci
Novo city, 1918, J. Rick s.n. (SP33928 as P. egregius Rick nom. in herb.).
FIGURE 3. Pluteus cervinus sensu Rick (PACA14519). a. Basidiospores; b. Pleurocystidia.
Scale bars = 10 µm.
Capítulo I - 20
Capítulo I - 21
COMMENTS — The re-examination of the collections identified by Rick as
P. cervinus shows that they actually represent specimens of P. xylophilus.
Although it was not possible to find any cheilocystidia, the characteristic
pleurocystidia in combination with the size and shape of the basidiospores
and the macroscopic aspect of the pileus are distinctive enough to identify
ick’s collections. Pluteus xylophillus can be easily recognized by its large
pileus, the presence of three types of pleurocystidia and the broadly ellipsoid
to elongate basidiospores (Singer 1959, Menolli et al. 2010).
There is much discussion about the identity and circumscription of
P. cervinus (Schaeff.) P. Kumm., and it is likely restricted to the northern
hemisphere, as shown in the phylogenetic analyses published by Justo et al.
(2011b). As suggested by Singer (1959), almost all collections mentioned
from Brazil as P. cervinus are probably P. xylophilus, a species that is well
known in South America and it forms a distinct clade in sect. Pluteus (Justo
et al. 2011b).
Rick (1907) also recorded P. cervinus var. patricius; however, he did not
mention the collection studied. No sample collected by Rick under this name
was found in the herbaria, but it most likely also represents P. xylophilus.
Another collection (PACA14529) of P. cervinus recorded by Rick (1961) is
deposited at PACA as P. wehlianus (see data under this species); however,
we did study PACA14526, which was named as P. cervinus and, due to a
typographical error, may represent the published PACA14529.
Pluteus cervinus var. griseoviridis Henn., Hedwigia 43: 204. 1904.
FIGURES 4, 5
— Brazil, São Paulo State, São Paulo city, Serra da
Cantareira, 1903, A. Puttemans 871 (Holotype – SP141797).
SPECIMEN EXAMINED
ORIGINAL DESCRIPTION — “Pileo carnoso, convexo explanato, centro obtuso,
laevi glabro, subglutinosulo, margine striatulo, griseo-viridi, 5–7 cm diam.;
stipite solido, cylindraceo, pallido, laevi, glabro, 4–7 × 3–5 cm, basi
incrassato; lamellis liberis, confertis, ventricosis, ca. 3–4 mm latis, flavidoincarnatis; sporis ellipsoideis, 1-guttulatis, carneis, 4–5 × 4 µ; cystidiis
lageniformibus, apice 2-vel 3 dentato-hamatis, 40–60 ×15–20 µ.”
MICROCHARACTERS — BASIDIOSPORES [100/5/5] 5.0–7.5(–8.7) × (3.1–)3.7–
5.0 µm [Q = 1.24–1.70(–2.03); Qm = 1.57; Lm = 6.3 µm; Wm = 4.1 µm],
broadly ellipsoid, ellipsoid to elongate, rarely cylindrical, inamyloid, hyaline,
smooth, thick-walled, guttulate. P LEUROCYSTIDIA 50–95 × 10.0–24 µm,
fusoid-ventricose, colorless, hyaline, mostly of the Cervinus-type with four to
six apical or lateral prongs, or like those found in P. harrisii that are usually
without or with two short lateral prongs, sometimes with lateral secondary
spinules such as those of P. spinulosus Murrill, thick-walled, abundant to
very abundant. CHEILOCYSTIDIA not recovered in most collections but a
clavate form (39 × 15.0 µm) was found in BPI770905. PILEIPELLIS a cutis.
CLAMP CONNECTIONS absent.
ADDITIONAL SPECIMENS EXAMINED — Brazil, Rio Grande do Sul State, Arroio
do Meio city, 1920, J. Rick s.n. (SP33926 as P. cervinus); 1920, J. Rick s.n.
(BPI770858 as P. cervinus); 1920, J. Rick s.n. (BPI770893 as P. princeps Rick
nom. in herb.); 1920, J. Rick s.n. (BPI770905 as P. umbrosus); São Leopoldo
city, 1929, leg. Steffen, det. J. Rick (PACA14517 as P. dissimilis Rick nom. in
herb.).
FIGURE 4. Pluteus cervinus var. griseoviridis (holotype – SP141797). a. Basidiospores; b.
Pleurocystidia. Scale bars = 10 µm.
COMMENTS — The re-examination of the holotype of P. cervinus var.
griseoviridis, along with some unpublished collections identified by Rick as
P. cervinus and P. umbrosus, and others named as “P. dissimilis” nom. in
herb. and “P. princeps” nom. in herb. showed that they share some
morphological features with P. angustisporus described from Bolivia (Singer
1959).
Pluteus angustisporus is morphologically close to P. xylophilus and
P. harrisii, from which it differs mainly by the narrow basidiospores and the
absence of pleurocystidia of the Magnus-type as found in P. xylophilus.
Capítulo I - 22
Capítulo I - 23
Although the collections we examined have large pilei, such as those
observed in P. xylophilus (up to 110 mm according to Menolli et al. 2010)
and much bigger than those reported for P. angustisporus (up to 25 mm
according to Singer 1959), the pleurocystidia of the Magnus-type were not
found in any of the collections studied. In addition, Singer (1959) also
reported for P. angustisporus the occurrence of pleurocystidia with
polymorphic apices like those observed (Fig. 5). However, due to the
preservation state of the exsiccatae, it was not possible to confirm the
presence of polymorphic cheilocystidia and pileipellis elements as reported
by Singer (1959).
It is not possible to confirm the actual identity of these collections from
these observations, but based on the recovered data, these collections are
clearly close to or identically with either P. angustisporus or P. xylophilus,
and we tentatively conclude that the former is the most appropriate name for
them.
It is important to note that although P. cervinus var. griseoviridis was
described before P. angustisporus, if they are the same species, the first name
does not have priority at species rank.
FIGURE 5. Pluteus cervinus sensu Rick (BPI770858). a. Basidiospores; b. Pleurocystidia. Scale
bars = 10 µm.
“Pluteus dissimilis Rick”, nom. in herb.
FIGURE 6
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, São Leopoldo city,
1929, leg. Steffen, det. J. Rick (PACA14517).
Capítulo I - 24
MICROCHARACTERS AND COMMENTS — See under P. cervinus var.
griseoviridis, with which it shares the same features. It is also considered a
probable P. angustisporus.
FIGURE 6. Pleurocystidia of Pluteus dissimilis nom. in herb. (PACA14517). Scale bar = 10 µm.
“Pluteus egregius Rick”, nom. in herb.
FIGURE 7
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Pareci Novo city,
1918, J. Rick s.n. (SP33928).
MICROCHARACTERS AND COMMENTS — See under P. cervinus, with which it
shares the same features. It is also considered P. xylophilus.
Pluteus exiguus var. venosus Rick, Iheringia Bot. 8: 417. 1961. as ‘venosa’.
nom. inv.
FIGURE 8
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Marcelino Ramos
city, 1936, J. Rick s.n. (PACA14518 as P. exiguus var. ‘venosa’).
ORIGINAL DESCRIPTION —“Pileo brunneo, venoso, lamellis mediis, densis,
stipite vitreo. Sporis 4–5 µm. Ad ligna.”
MICROCHARACTERS — BASIDIOSPORES [20/1/1] 5.6–7.5 × (4.6–)5.0–6.2 µm
[Q = (1.09–)1.12–1.24); Qm = 1.20; Lm = 6.3 µm; Wm = 5.3 µm],
FIGURE 7. Pleurocystidia of Pluteus egregius nom. in herb. (SP33928). Scale bar = 10 µm.
subglobose to broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled,
guttulate. PILEIPELLIS a hymeniderm, composed of subglobose to
spheropedunculate cells, 26–37 × 18.7–26.5 µm, almost non-pedicellate or
with a short pedicel (1.2–3.7 µm long), thin-walled, with brown dissolved
content. PLEURO- AND CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS
absent.
COMMENTS — Although the package of the collection PACA14518 is named
as P. exiguus var. ‘venosa’, it was publis ed by ick (19 1 under t e
taxonomic entry of P. exiguus and not under P. exiguus var. ‘venosa’. Thus,
because the holotype was not clearly designated in the protologue, the name
can be considered a nomen invalidum.
Despite the nomenclatural problems, the occurrence of P. exiguus (Pat.)
Sacc. or any variety of this species in Brazil seems unlikely. It is important to
consider that the description of P. exiguus available in Rick (1961) is almost
identical to that provided by Saccardo (1887) and that no more material under
this name was located in the herbaria, making it difficult to confirm its
occurrence in Brazil. In addition, as far as we know, the occurrence of
Capítulo I - 25
Capítulo I - 26
P. exiguus appears to be restricted to Europe, where it has been barely
documented (Kühner & Romagnesi 1956, Orton 1986, Justo & Castro 2007a,
Ripková 2009).
The proposal of the new variety by Rick (1961) seems inappropriate
because collection PACA14518 has a hymenidermal pileipellis typical of
sect. Celluloderma rather than a pileipellis typical of sect. Hispidoderma
Fayod, under which P. exiguus is classified. The scarce morphological data
obtained from ick’s collection are insufficient to designate a correct name
for it, but we agree with Singer (1959) that Rick most likely had
P. jamaicensis, which is characterized by non-globose basidiospores [5.0–
6.8(–7.5) × 4.5–6.2 µm] and pilleipellis cells (21–60 × 20–45 µm) with
brown dissolved content (Singer 1956, 1959; Smith & Stuntz 1958; Horak
1964; Pegler 1983; Banerjee & Sundberg 1993; Menolli et al. 2010).
FIGURE 8. Pluteus exiguus var. venosus (PACA14518). a. Basidiospores; b. Pileipellis cells.
“Pluteus fuscidulus Fr.”, nom. in herb.
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, São Salvador city, 22
Jul 1944, J. Rick s.n. (PACA20926).
COMMENTS — This collection is too poor in condition for study of any
microstructures or to gain insights about its identification. Moreover, there is
no Pluteus (nor Agaricus) named by Fries as ‘fuscidulus’. Because the
collection is unfit for study and the name not attributable to Fries, we have
nothing to add to this record.
“Pluteus granulatus Bres.” sensu Rick, Iheringia Bot. 8: 418. 1961.
FIGURE 9A
1929, J. Rick s.n. (PACA14523*).
MICROCHARACTERS — BASIDIOSPORES [20/1/1] 6.2–7.5 × 5.0–6.2 µm (Q =
1.21–1.50; Qm = 1.28; Lm = 7.2 µm; Wm = 5.7 µm), preponderantly broadly
ellipsoid or sometimes ellipsoid, inamyloid, hyaline, smooth, thick-walled,
guttulate. PILEIPELLIS composed of hyphal elements typical of a cutis but it is
too difficult to be recovered. PLEURO- AND CHEILOCYSTIDIA not recovered.
CLAMP CONNECTIONS not observed.
COMMENTS — Pluteus granulatus is a rarely recorded mushroom that seems
to be restricted to Europe (Saccardo 1887, Kühner & Romagnesi 1956, Orton
1986) and it is considered a synonym of P. plautus (Weinm.) Gillet – in a
broad sense (Vellinga & Schreurs 1985, Legon & Henrici 2011).
Singer & Digilio (1952) recorded P. plautus from Argentina, but later
Singer (1959) re-identified the collection as P. argentinensis Singer.
According to Singer (1959), Rick’s collection was misidentified, and our reexamination suggests that it most likely represents some species with cutislike pileipellis characterized by an umber fibrillose pileus and broadly
ellipsoid to ellipsoid basidiospores. It is possible that Rick had
P. argentinensis, P. fastigiatus Singer, P. fuliginosus Murrill or most likely
P. glaucotinctus that was recently recorded from Brazil (Justo et al. 2011ab).
“Pluteus hispidulus Fr.” sensu Rick, Iheringia Bot. 8: 419. 1961. FIGURE 9B, C
Rick s.n. (PACA14528*); 1932, Rick s.n. (FH00301668); São Salvador city,
1943, Rick s.n. (PACA14532*).
1.16–1.50; Qm = 1.30; Lm = 7.8 µm; Wm = 6.0 µm), broadly ellipsoid to
ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS
and HYMENIAL STRUCTURES not recovered. CLAMP CONNECTIONS not
observed.
ADDITIONAL SPECIMEN EXAMINED — Brazil, J. Rick s.n. (BPI839783).
COMMENTS — Pluteus hispidulus (Fr.) Gillet was described from Europe and
has not yet been definitively recorded from South America. Because the data
recovered are not sufficient to allow the renaming of Rick’s collections, it is
not possible to determine what species Rick had collected.
Singer (1959) suggested that this might be a species of sect.
Hispidoderma (stirps Fuliginosus). Among the probable species, it might be
P. yungensis that has the biggest basidiospores, which could be comparable
in size to those observed in Rick’s collections. Modern collections with these
characteristics are needed to clarify whether this species really occurs in
Brazil. Another unpublished collection (BPI839783), named by Rick as
P. hispidulus, was revised, but it actually represents an Entoloma s.l.
Capítulo I - 27
Capítulo I - 28
FIGURE 9. Basidiospores. a. Pluteus granulatus sensu Rick (PACA14523); b–c. Pluteus
hispidulus sensu Rick; b. PACA14528; c. PACA14532. Scale bars = 10 µm.
“Pluteus leoninus Schaeff.” sensu Rick, Iheringia Bot. 8: 417. 1961.
FIGURE 10
SPECIMEN EXAMINED — Brazil, Rio Grande Sul State, São Leopoldo city,
1932, J. Rick s.n. (FH00301669).
MICROCHARACTERS — BASIDIOSPORES [20/1/1] 5.0–6.2(–7.5) × 3.7–4.6
(–5.0) µm [Q = (1.09–)1.22–1.51(–1.68); Qm = 1.37; Lm = 5.7 µm; Wm =
4.2 µm), broadly ellipsoid to ellipsoid, rarely subglobose or elongate,
inamyloid, hyaline, smooth, thick-walled, guttulate. PLEUROCYSTIDIA 52–87
× 12.5–26 µm, fusoid-ventricose to lageniform, usually with a rounded apex,
metuloidal with thickened wall and lacking any apical or lateral horns,
frequently with pale straw or colorless internal condensed or dissolved
content. CHEILOCYSTIDIA not recovered. PILEIPELLIS difficult to recover but
apparently a cutis composed of hyphae with pale brown pigment. CLAMP
CONNECTIONS absent.
COMMENTS — Pluteus leoninus (Schaeff.) P. Kumm. belongs to sect.
Hispidoderma, and it is apparently restricted to the northern hemisphere. The
species is characterized by golden yellow basidiomata and by thin-walled
pleurocystidia with little excrescences at the apex (Singer 1956, Vellinga &
Schreurs 1985, Orton 1986, Takehashi & Kasuya 2007). However,
P. leoninus sensu Rick has thick-walled pleurocystidia. Considering other
species with yellowish basidiomata and non-horned metuloidal
pleurocystidia, it is likely that Rick had P. conizatus or P. amphicystis Singer,
which differ from each other in pileipellis structure and the absence of
metuloidal caulocystidia in P. conizatus (Singer 1956, 1959; Pegler 1983,
1986; Pradeep et al. 2002 . ecause t e pileipellis in ick’s collection was
not well recovered, we verified that the caulocystidia are absent in this
collection, and it is probable that Rick actually had P. conizatus instead of
P. leoninus.
FIGURE 10. Pluteus leoninus sensu Rick (FH00301669). a. Basidiospores; b. Pleurocystidia.
Pluteus leptonia Rick, Lilloa 3: 445. 1938.
1936, J. Rick s.n. (Holotype – PACA14525*).
ORIGINAL DESCRIPTION — “Pileo conico, demum expanso-fisso, carnoso, 3–4
cm. lato, subumbonato, laevi, minute ruguloso, exstrio; lamellis aperte
liberis, densis, subventricosis, evidenter erosis carneis; stipite usque 1 dm.
Longo 5 mm lato, firmo, demum cavo, nudo, striato, cartilagineo; sporis
angulatis 7–10 micr. apiculatis. Omnibus partibus pulcherrime lazulinus aut
coeruleus exceptis lamellis carneis e carne carnea. Odore et sapore nullo. In
terra inter Bambusas. Pluteus phaeus Sacc. XVI et sine dubio proximus nisi
identicus, sed mea species non crescit in ligno, sed in terra.”
COMMENTS — The type of P. leptonia is not a Pluteus; it is actually a
member of Entolomataceae. Its basidiospores are predominantly
isodiametric, with 5–7 angles. Singer (1956, 1959) mentioned the presence of
typical Rhodophyllus basidiospores, and e suggested t at ick’s collection
could be Nolanea howellii Peck or another allied species.
Capítulo I - 29
Capítulo I - 30
“Pluteus longistriatus Peck” sensu Rick, nom. in herb.
SPECIMEN EXAMINED — Brazil, Rio Grande Sul State, São Leopoldo city,
1932, J. Rick s.n. (FH00301670).
1.07–1.24; Qm = 1.19; Lm = 7.0 µm; Wm = 5.9 µm), subglobose to broadly
ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS
and HYMENIAL STRUCTURES not recovered. CLAMP CONNECTIONS not
observed.
COMMENTS — Pluteus longistriatus (Peck) Peck is common in subtropical
and temperate areas of North and South America and it is usually recognized
by a pileus with a deeply sulcate-striate margin and a hymenidermal
pileipellis with heteromorphic elements (Singer 1959, Menolli & Capelari
2010). Considering the macroscopic appearance of FH00301670, it is
possible to suppose that it had a sulcate-striate pileus when it was fresh, and
it is important to consider that the basidiospore size and shape are in
accordance with those usually reported for P. longistriatus (Singer 1959,
Menolli & Capelari 2010 . ased on t ese data, we suspect t at ick’s
collection is actually P. longistriatus, but we cannot confirm the
identification and the occurrence of this species from Rio Grande do Sul
State with certainty because the pileipellis and the hymenial structures were
not recovered from this collection.
“Pluteus murinus Bres.” sensu Rick, nom. in herb.
FIGURE 11
SPECIMEN EXAMINED — Brazil, J. Rick s.n. (BPI770882).
MICROCHARACTERS — BASIDIOSPORES [40/2/2] (6.2–)7.5–8.7 × (5.0–)6.2
–7.5 µm [Q = (1.00–)1.16–1.24(–1.60); Qm = 1.19; Lm = 7.6 µm; Wm = 6.4
µm), preponderantly broadly ellipsoid or sometimes globose or ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. PLEUROCYSTIDIA 51
–69(–86) × 10.0–22.5 µm, elongate-clavate or slightly ventricose, thin- to
relatively thick-walled, sometimes with a colorless internal condensed
content. CHEILOCYSTIDIA 50–66 × 12.5–15 µm, clavate with a welldeveloped pedicel at the base, thin-walled, colorless and hyaline. PILEIPELLIS
a cutis composed of hyphal elements with a rounded apex and brownish
dissolved pigment. CLAMP CONNECTIONS not observed.
ADDITIONAL SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, São
Salvador city, 1944, J. Rick s.n. (PACA20770 as P. sensitivus var.
macrospora).
Capítulo I - 31
COMMENTS — Pluteus murinus is apparently restricted to Europe (Singer
1956, Orton 1986, Banerjee & Sundberg 1993), and it is also considered a
synonym of P. ephebeus (Fr.) Gillet (Vellinga & Schreus 1985). Considering
the data recovered from Rick’s collections, we see that it shares the main
characteristics found in P. albostipitatus, a widely distributed and commonly
cited species with some synonyms and wide morphological variation, mainly
regarding the presence of fibrils on the pileus, the wall thickness in the
pleurocystidia and the size and shape of the basidiospores (Menolli et al.
2010; Justo et al. 2011b).
FIGURE 11. Pluteus murinus sensu Rick (BPI770882). a. Basidiospores; b. Pleurocystidia;
c. Cheilocystidia. Scale bars = 10 µm.
“Pluteus nanus Pers.” sensu ick, I eringia ot. 8: 418. 19 1.
FIGURE 12
SPECIMENS EXAMINED — Brazil, Rio Grande do Sul State, Marcelino Ramos
city, 1936, Rick s.n. (FH00301671); São Salvador city, 29 Feb. 1944, Rick s.n.
(PACA22618*).
MICROCHARACTERS — BASIDIOSPORES [40/2/2] 5.0–7.5 × 5.0–6.2(–7.5) µm
[Q = 1.00–1.12(–1.24); Qm = 1.08; Lm = 6.3 µm; Wm = 5.9 µm], globose or
occasionally broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled,
guttulate. PILEIPELLIS a hymeniderm composed of subglobose cells, 20–59 ×
15.0–34 µm, with a short to moderately long pedicel (1.2–10.0 µm long),
thin-walled, with brownish condensed or dissolved content. PLEURO- AND
CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS not observed.
COMMENTS — Pluteus nanus (Pers.) P. Kumm. is a species with many
interpretations, and it seems to be restricted to Europe (Vellinga & Schreurs
1985, Orton 1986, Justo et al. 2007, Justo et al. 2011b). Although there is a
suspect record from North America (Homola 1972), it remains unclear if
P. nanus is really found in the USA according to any of these interpretations
(Minnis & Sundberg 2010). Singer (1959) considered Rick’s identification to
be dubious. Considering the rugose-venose pileus, the globose basidiospores
and the pileipellis cells with brownish condensed or dissolved content, it is
possible that Rick had P. pulverulentus or most likely P. sapiicola, a species
known to occur in Brazil (Menolli, de Meijer & Capelari, unpublished data).
However, other species with pigmented cystidia, such as P. beniensis Singer,
P. eliae Singer, P. rimosoaffinis, and P. riograndensis Singer, could also be
considered.
Rick (1938, 1961) also recorded P. nanus var. podospileus, but he did not
mention the material studied. Because there are no vouchers concerning the
record of P. nanus var. podospileus from Brazil and because no other
collections under this name were located, nothing can be added to this record.
FIGURE 12. Pluteus nanus sensu Rick. a–b. FH00301671. a. Basidiospores; b. Pileipellis cells.
c–d. PACA22618. c. Basidiospores; d. Pileipellis cells. Scale bars = 10 µm.
Capítulo I - 32
“Pluteus pellitus Pers.” sensu Rick, Iheringia Bot. 8: 418. 1961.
Capítulo I - 33
FIGURE 13
Rick s.n. (PACA14524*); 1931, Rick s.n. (FH00301673); 1932, Rick s.n.
(PACA14530*); Sep. 1933, Rick s.n. (FH00301672).
MICROCHARACTERS — BASIDIOSPORES [80/4/4] (5.6–)6.2–7.5(–8.7) × (3.7–)
5.0–6.2 µm [Q = 1.21–1.51(–1.68); Qm = 1.28; Lm = 6.7 µm; Wm = 5.3
µm], preponderantly broadly ellipsoid or sometimes ellipsoid, rarely
elongate,
inamyloid,
hyaline,
smooth,
thick-walled,
guttulate.
PLEUROCYSTIDIA 61–78 × 16.2–18.7 µm, slightly ventricose to lageniform,
with an amorphous pale straw content, pigmentation usually concentrated at
apex and base, thin-walled (only recovered in FH00301673).
CHEILOCYSTIDIA and PILEIPELLIS not recovered. CLAMP CONNECTIONS not
observed.
FIGURE 13. Pluteus pellitus sensu Rick. a–c. Basidiospores. d. Pleurocystidia. a. PACA14524;
b. PACA14530; c–d. FH00301673. Scale bars = 10 µm.
COMMENTS — Although there are many concepts of P. pellitus (Pers.)
P. Kumm., Justo and Castro (2007b) and Justo et al. (2011b) accepted it as a
species that is apparently restricted to Europe and is characterized mainly by
the white basidiomata, a habitat on angiosperm wood, basidiospores of 5.0
–7.5 × 3.5–5.0 µm, and the presence of clamp connections. Singer (1959)
mentioned that P. pellitus sensu ick “is obviously P. viscidulus”, alt oug
e did not mention ick’s collection in t e ‘e amined material’ for t is
species. Pluteus viscidulus Singer is another species with white basidiomata
but without clamp connections. It was described by Singer (1959) from
Argentina and it was also recorded from Brazil (from the same State from
w ic
ick’s material was collected . Singer (1959 also recorded
P. viscidulus from the U.S.A. (Florida), although he mentioned that the North
American collection had bigger basidiospores and that this character can be
used to separate it from P. viscidulus.
The re-examination of the Brazilian collection determined by Singer as
P. viscidulus (Fig. 14) [Brazil, Rio Grande do Sul State, São Salvador city,
9 Nov. 1951, Singer B111 (LIL)] showed that it has broadly ellipsoidal to
elongate basidiospores, [20/1/1] (5.6–)6.2(–7.5) × 3.7–5.0 µm [Q = 1.24
–1.68; Qm = 1.35; Lm = 6.3 µm; Wm = 4.7 µm], fusoid-ventricose
pleurocystidia (69–83 × 11.2–25 µm) that have thick walls and apices with
two to four lateral prongs, clavate cheilocystidia (37–61 × 12–15 µm),
pileipellis that seems to be a cutis, and lack of clamp connections. Similar
features were also found by Justo and Castro (2007b) in the isotype of
P. viscidulus. Apparently, up until that point, P. viscidulus had represented an
authentic South American species with white basidiomata and without
clamp-connections; however, based on molecular analyses, Justo et al.
(2011b) considered P. viscidulus to fall within the molecular variation of
P. petasatus, and they can consequently be treated as synonyms.
Thus, although the occurrence of the white form of P. petasatus
(= P. viscidulus can be confirmed from razil based on Singer’s collection,
the morphological diagnostic characteristics for it were not observed in
ick’s collections, or at least for t e unpublis ed collection FH00301 73 t at
has the thin-walled and pigmented pleurocystidia instead of those observed in
P. pellitus and P. petasatus or allied species, which have the horned
pleurocystidia that are typical of sect. Pluteus. In addition, according to notes
for FH00301673, Rick denoted it as P. pellitus ‘videtur’, w ich indicates that
although it resembles P. pellitus, another allied species can also be
considered.
Because there are no notes about the pileus color of FH00301673, it is
difficult to suggest a name for it. There are no previously published species
that combine broadly ellipsoid basidiospores and pigmented pleurocystidia
Capítulo I - 34
FIGURE 14. Pluteus viscidulus (Singer B111 – LIL). a. Basidiospores; b. Pleurocystidia;
with a whitish pileus. However, because there is no mention whether the
pileus is actually whitish, if we consider only the species of sect.
Hispidoderma with pigmented pleurocystidia, it could be any species from
stirps Aethalus, Circumscissus or Plautus, according to Singer’s classification
(Singer 1959).
Capítulo I - 35
Capítulo I - 36
Although no reliable conclusion can be made about the identity of Rick’s
collections, we suggest that they are not P. pellitus. We should not disregard
the possibility that the collections (PACA14524, PACA14530) published in
Rick (1961) could be P. petasatus, and FH00301673 is most likely a species
of sect. Hispidoderma.
“Pluteus princeps Rick”, nom. in herb.
FIGURE 15
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Arroio do Meio city,
1920, J. Rick s.n. (BPI770893).
MICROCHARACTERS AND COMMENTS — See under P. cervinus var.
griseoviridis, with which it shares the same features. It is also considered a
probable P. angustisporus.
FIGURE 15. Pluteus princeps nom. in herb. (BPI770893) a. Basidiospores; b. Pleurocystidia.
“Pluteus phlebophorus Dittm.” sensu Rick, Iheringia Bot. 8: 419. 1961.
FIGURE 16
1934, leg. Steffen, det. J. Rick (PACA14520*); São Salvador city, 28 Jan 1944,
J. Rick s.n. (PACA20895).
MICROCHARACTERS (PACA14520) — BASIDIOSPORES [20/1/1] 5.0(–6.2) ×
3.7(–5.0) µm [Q = (1.24–)1.35(–1.68); Qm = 1.39; Lm = 5.2 µm; Wm = 3.8
µm], preponderantly ellipsoid or sometimes broadly ellipsoid or elongate,
inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS a
hymeniderm composed of subglobose cells, 24–36 × 18.7–22 µm, with a
short pedicel (2.5–3.7 µm long), thin-walled, with brownish condensed
content. PLEURO- AND CHEILOCYSTIDIA not recovered. CLAMP CONNECTIONS
not observed.
MICROCHARACTERS (PACA20895) — BASIDIOSPORES [20/1/1] 5.0–6.2 ×
5.0–6.2 µm (Qm = 1.00; Lm = 5.7 µm; Wm = 5.7 µm), globose, inamyloid,
hyaline, smooth, thick-walled, guttulate. PILEIPELLIS a hymeniderm
composed of subglobose cells, 31–36 × 16.2–25 µm, with a short to
moderately long pedicel (2.5–12.5 µm long), thin-walled, with brownish
condensed or dissolved content. PLEURO- AND CHEILOCYSTIDIA not
recovered. CLAMP CONNECTIONS not observed.
FIGURE 16. Pluteus phlebophorus sensu Rick.
a–b. PACA14520. a. Basidiospores;
b. Pileipellis cells. c–d. PACA20895. c. Basidiospores; d. Pileipellis cells. Scale bars = 10 µm.
Capítulo I - 37
Capítulo I - 38
COMMENTS — These two collections studied by Rick have different
micromorphological characteristics and represent two species, although they
were both listed as P. phlebophorus.
Singer (1959) suggested that P. phlebophorus sensu Rick is likely
P. jamaicensis, which is characterized by non-globose basidiospores of 5.0
–6.8(–7.5) × 4.5–6.2 µm (Singer 1956, 1959; Smith & Stuntz 1958; Horak
1964; Pegler 1983; Menolli et al. 2010), differing in size and shape from
t ose observed in bot ick’s collections. PACA14520 most likely represents
P. tucumanus, and it shares characteristics with PACA14522, named by Rick
as P. brunneopictus (see discussion under P. brunneopictus). Additionally,
PACA20895 is most likely P. sapiicola, similar to PACA22618 named by
Rick as P. nanus (see discussion under P. nanus).
“Pluteus sensitivus var. macrospora Rick”, nom. in herb.
FIGURE 17
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, São Salvador city,
1944, J. Rick s.n. (PACA20770).
MICROCHARACTERS — See under P. murinus, with which it shares the same
features.
COMMENTS — It is important to explain that Rick (1930) described
P. sensitivus without mentioning the holotype, but later Rick (1961)
mentioned PACA20770 as the material studied for this species. However,
according to the exsiccata notes, the collection PACA20770 is an
unpublished variety, viz. P. sensitivus var. macrospora, and represents the
only collection found for the epithet ‘sensitivus’. Although PACA20770 was
published under P. sensitivus (Rick 1961), it obviously does not represent the
holotype because P. sensitivus was described in 1930 and this material was
collected in 1944. Since there are no more data about the holotype of
P. sensitivus, and because no other collections under this name were located,
it can be considered a nomen dubium. Furthermore and based on the data
recovered from the specimen named as P. sensitivus var. macrospora, we
suggest that it may actually be P. albostipitatus (see discussion under
P. murinus).
Pluteus straminellus Rick, Iheringia Bot. 8: 417. 1961.
1930, J. Rick s.n. (Holotype – PACA15531*).
FIGURE 17. Pluteus sensitivus var. macrospora nom. in herb. (PACA20770) a. Basidiospores;
b. Pleurocystidia. Scale bars = 10 µm.
ORIGINAL DESCRIPTION — “Pileo stramineo, innato-squamuloso, centro
lurido, stipite brevi, aquoso; lamellis densis, arcuatis, carneolis, liberis.
Sporis 4 × 3 µm. Ad lignum.”
MICROCHARACTERS — BASIDIOSPORES [20/1/1] 3.7(–4.4) × 2.5(–3.1) µm
[Q = (1.42–)1.48; Qm = 1.48; Lm = 3.7 µm; Wm = 2.5 µm), ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS and
HYMENIAL STRUCTURES not recovered. CLAMP CONNECTIONS not observed.
COMMENTS — Based only on the size and shape of basidiospores, it is not
possible to establish P. straminellus as a sufficiently well-known species,
leading us to treat it as a nomen dubium. If there were data about the
pileipellis and hymenial structures and more details about the pileus color, it
would be possible to suggest a morphological relationship between
P. straminellus and other species with small basidiospores, such as
P. microsporus (Dennis) Singer, P. ugandensis Pegler or P. unakensis
Murrill.
Capítulo I - 39
“Pluteus umbrosus Pers.” sensu Rick, Iheringia Bot. 8: 418. 1961.
Capítulo I - 40
FIGURE 18
SPECIMEN EXAMINED — Brazil, Rio Grande do Sul State, Arroio do Meio city,
1920, J. Rick s.n. (BPI770905).
MICROCHARACTERS — See under P. cervinus var. griseoviridis, with which it
shares the same features.
COMMENTS — It is important to note that the name attributed by Rick (1919,
1938, 1961) is obviously inaccurate because P. umbrosus (Pers.) P. Kumm. is
a species of sect. Hispidoderma (Singer 1956, Vellinga & Schreurs 1985,
Orton 1986) and the collection studied here has horned pleurocystidia that are
typical of sect. Pluteus. Based on the morphological data recovered, this
Rick’s collection is also considered a probable P. angustisporus (see
discussion under P. cervinus var. griseoviridis).
FIGURE 18. Pluteus umbrosus sensu Rick (BPI770905). a. Basidiospores; b. Pleurocystidia;
Pluteus velatus Rick, Iheringia Bot. 8: 417. 1961.
Capítulo I - 41
FIGURE 19
1939, J. Rick s.n. (Holotype – PACA14527* as P. velutinus).
ORIGINAL DESCRIPTION — “Campanulatus, 3 cm latus, griseo-brunneus,
membranaceus, squarrosulus ad marginem, secus nudus, fere usque ad
centrum leviter striatus; lamellis liberis, ventricosis, densis, albis, margine
rubro. Basidiis brevibus, cystidiis paucis, latis; stipite 6 cm alto, 3 cm lato,
subtus crassiore, albo-lanoso, ceterum nudo, striatulo, pallido, velo
arachnoideo. Sporis 6–10 × 5–10 µm, polygonalibus, rubris. Ad lignum
putridum.”
FIGURE 19. Pluteus velatus (holotype – PACA14527) a. Basidiospores; b. Pleurocystidia. Scale
bars = 10 µm.
MICROCHARACTERS — BASIDIOSPORES [20/1/1] (6.2–)7.5–8.7(–10.0) × 5.0
–6.2(–7.5) µm [Q = (1.16–)1.21–1.50(–1.61); Qm = 1.32; Lm = 7.9 µm;
Wm = 6.0 µm), broadly ellipsoid to ellipsoid or rarely almost elongate,
inamyloid, hyaline, smooth, thick-walled, guttulate. PILEIPELLIS composed of
hyphal elements typical of a cutis but it is too difficult to be recovered.
PLEUROCYSTIDIA 56–69(–87) × (13.7–)18.7–24(–30) µm, clavate or slightly
ventricose, thin-walled. CHEILOCYSTIDIA not recovered. CLAMP
CONNECTIONS not observed.
COMMENTS — Singer (1954, 1959 recognized ick’s collection as a
representative of P. fibulatus Singer described from Argentina (Singer &
Digilio 1952). It is not clear, however, whether Singer actually analyzed this
collection; nevertheless, he also mentioned “t at t e indication of a veil in t e
unpublis ed notes must be due to an error”. Pluteus velatus was published in
the posthumous paper of Rick (1961), and according to notes of Singer
(1954, 1959) it should be considered a synonym of P. fibulatus. However, the
re-examination of the types of P. fibulatus and P. velatus confirms that they
represent different species.
Pluteus fibulatus is a typical species of sect. Pluteus, characterized by a
fibrillose and deep-colored pileus, the presence of clamp-connections and the
metuloidal and horned pleurocystidia (Singer 1959). The re-examination of
the isotype of P. fibulatus (Fig. 20) [Argentina, Prov. Tucuman, Capital, 22
Feb. 1949, Singer T141 (MICH)] showed that it has globose to broadly
ellipsoidal or rarely ellipsoidal basidiospores, [20/1/1] (5.6–)6.2–7.5(–8.7)
× (5.0–)5.6–6.8(–7.5) µm [Q = 1.00–1.24(–1.40); Qm = 1.16; Lm = 7.1 µm;
Wm = 6.1 µm]; fusoid-ventricose pleurocystidia (65–88 × 17.5–28.7 µm)
that have thickened or moderately thickened walls and apices provided with
two to four apical prongs (or sometimes without prongs and with an acute or
rounded apex); clavate cheilocystidia (44 × 13.7 µm); pileipellis that seems
to be a cutis but that is difficult to recover, and clamp connections that are
present but difficult to detect. Similar features were also found in the
collection of P. fibulatus (Fig. 21) recorded by Pegler (1997) from São Paulo
State, Brazil [Brazil, São Paulo State, Mogi Guaçu city, 30 Jan. 1987, Pegler
et al. 3899, K(M)41525]. However, globose basidiospores and metuloidal or
horned cystidia were not found after the re-examination of the holotype of
P. velatus. All the pleurocystidia observed in P. velatus had thin walls, which
indicates that it is likely to be a member of sect. Hispidoderma rather than
P. fibulatus or any other species of sect. Pluteus. Thus, we can confirm the
occurrence of P. fibulatus in razil, but t e recovered data from ick’s
collection are not sufficient to confirm the identity of P. velatus, and it
remains a nomen dubium. Taking into consideration the macroscopic
characteristics described by Rick (1961), such as the pileus color and the
Capítulo I - 42
presence of lamellae with colored edges (‘margine rubro’), as well as the size
and shape of the basidiospores and pleurocystidia described herein, it is likely
that the material collected by Rick is P. fernandezianus Singer from Chile
(Singer 1959).
FIGURE 20. Pluteus fibulatus (isotype – Singer T141, MICH). a. Basidiospores;
b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 µm.
“Pluteus wehlianus Müller” sensu Rick, nom. in herb.
1939, J. Rick s.n. (PACA14529).
MICROCHARACTERS — BASIDIOSPORES [4/1/1] 16.2–20 × 11.2–12.5 µm
(Q = 1.4–1.6; Qm = 1.51; Lm = 18.4 µm; Wm = 12.2 µm), ellipsoid with
germ pore. PILEIPELLIS and HYMENIAL STRUCTURES not recovered. CLAMP
CONNECTIONS not observed.
COMMENTS — This is an unpublished collection that obviously does not
represent a Pluteus species due to the large basidiospores with germ pore. It
is likely a species of Bolbitiaceae or Strophariaceae.
Capítulo I - 43
FIGURE 21. Pluteus fibulatus [Pegler et al. 3899 – K(M)41525]. a. Basidiospores;
b. Pleurocystidia; c. Cheilocystidia. Scale bars = 10 µm.
Discussion
Among the 32 Pluteus names attributed by Rick and Hennings, only one
collection published as P. cervinus could be assigned to the correct name
P. xylophilus. Three other unpublished collections named as P. egregius,
P. murinus and P. sensitivus var. macroscpora were correctly identified as
P. xylophilus and P. albostipitatus (the two latter), respectively. Materials of
ten taxa were not found and at least five of them could be considered nomen
dubium, besides P. straminellus and P. velatus that we also consider as
nomen dubium because based on the morphological data recovered from
Rick’s collections are not sufficient to confirm the identity of them.
From the remainder, we only suggested another name for the collections
of Rick and Hennings because it was not possible to recover complete
morphological data for a correct identification, although the data available
are sufficient to consider inappropriate the names originally attributed by
them. In addition, there is no sufficient material of P. fuscidulus (nom. in
herb.) for study, and finally P. leptonia is an Entolomataceae and
P. wehlianus (unpublished) belongs to Bolbitiaceae or Strophariaceae.
Capítulo I - 44
Acknowledgments
The authors thank Andrew M. Minnis and Else C. Vellinga for critical review and
for provided very insightful comments in the manuscript; all of the curators who
helped with this work, especially those from BPI, FH, K, LIL, MICH, PACA and SP
for the loan of collections; Klei R. Sousa for inking the illustrations; Fernanda
Karstedt for her assistance regarding the collections considered members of
Entolomataceae; the Graduate Program in “Biodiversidade Vegetal e Meio
Ambiente” for the financial support toward the costs involving this manuscript; and
the “Conselho Nacional de Desenvolvimento Científico e Tecnológico” (CNPq) for
the support and grant to the authors.
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Capítulo I - 47
Capítulo II - 48
CAPÍTULO II
Phylogeny of three widely distributed species of Pluteus (Pluteaceae, Agaricales):
taxonomic and biogeographic implications 
Nelson Menolli Jr., Alfredo Justo, Andrew M. Minnis, Pedro Arrillaga & Marina Capelari
Artigo submetido para publicação na revista Organisms Diversity & Evolution.
Capítulo II - 49
Phylogeny of three widely distributed species of Pluteus (Pluteaceae, Agaricales):
taxonomic and biogeographic implications
Nelson Menolli Jr.
Instituto Federal de Educação, Ciência e Tecnologia de São Paulo, Campus São Paulo, CCT / Biologia, Rua
Pedro Vicente 625, 01109-010 São Paulo, SP, Brazil
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902 São Paulo, SP, Brazil
Alfredo Justo
Biology Department, Clark University, 950 Main St., Worcester, MA 01610, USA
Andrew M. Minnis
Center for Forest Mycology Research, Northern Research Station, US Forest Service, One Gifford Pinochet Dr.,
Madison, WI 53726, USA
Pedro Arrillaga
Sociedad de Ciencias Aranzadi, Departamento de Micología, Alto de Zorroaga 11, San Sebastián 20014, Spain
Marina Capelari
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902 São Paulo, SP, Brazil
Corresponding author: Nelson Menolli Jr ([email protected])
Abstract:
In order to better understand the biogeography and global phylogeny of Pluteus and species delimitation
within the genus, we present a detailed study on the morphological and molecular (nrITS) variation of three
Pluteus species with putative intercontinental distributions, viz. Pluteus albostipitatus, P. glaucotinctus and
P. longistriatus. Pluteus albostipitatus is accepted here as a species with a relatively wide range of
morphological variation that is not correlated with variation at the molecular level. South American populations
seem to be more closely related to African populations than to North American populations. Pluteus
albostipitatus var. poliobasis, P. densifibrillosus, P. melanopotamicus and P. phaeoleucus are considered
synonyms of P. albostipitatus. Pluteus glaucotinctus is a pantropical species with specimens recorded from
countries in Africa, Asia, Europe and South America. The collections of P. glaucotinctus studied here likely
represent a species-complex with one of its representatives being widely distributed in South America, Asia and
Europe and two other representatives known only from single-collections in Africa. Collections of Pluteus
longistriatus from North and South America were found to be very similar in morphological and molecular
characters and therefore are considered as belonging to a single species that is widely distributed in subtropical
and temperate areas in the Americas.
Key words: Basidiomycota, Biogeography, Fungi, ITS, Species delimitation.
Capítulo II - 50
Introduction
Pluteus Fr. (Agaricales, Basidiomycota) comprises ca. 300 species with a circumglobal distribution (Kirk et al.
2008) and it is very common in forested areas from boreal to tropical regions. The genus is found primarily in
association with well-rotted wood and it is characterised by free lamellae, absence of volva, pinkish spore print,
inamyloid basidiospores and inverse hymenophoral trama (Singer 1986). Based on molecular data, it also
includes some annulate species (Minnis et al. 2006; Corriol and Moreau 2007; Vizzini and Ercole 2011). Recent
molecular studies (Menolli et al. 2010; Justo et al. 2011a; b) have more or less supported the traditional
subdivision of Pluteus into three sections, Pluteus, Celluloderma Fayod and Hispidoderma Fayod, while others
have increased the number of described species on the basis of morphological and molecular data (Rodríguez et
al. 2009; Rodríguez et al. 2010; Justo et al. 2011c; 2012; Pradeep et al. 2012).
Species recognition of agaric fungi has always been difficult and somewhat predisposed to subjectivity.
Besides morphological and molecular methods that are frequently used to distinguish mushroom species, the use
of sexual compatibility and interfertility as well as combining multiple methods with the different types of
evidence are alternative ways to recognize the limits of some taxa and to resolve taxonomic problems (Anderson
and Ullrich 1979; Chase and Ullrich 1990; Shimomura et al. 1992; Vilgalys and Sun 1994; Hibbett et al. 1995;
Zervakis and Balis 1996; Zervakis 1998; Petersen and Hughes 1999; Capelari and Fungaro 2003). Thus, recent
efforts to improve species recognition utilize a correlation between the three most common species concepts,
morphological, biological and phylogenetic (Taylor et al. 2000). However, the practical application of the
biological species concept is not feasible in Pluteus, due to the difficulty in obtaining basidiospore germination
under laboratory conditions (Banerjee and Sundberg 1993a).
Recent studies of macrofungi have also discussed their distributions and the biogeographical implications in
the context of morphological and molecular data (Hibbett 2001; Mueller et al. 2001; Oda and Tsuda 2004; Zhang
et al. 2004; Matheny and Bougher 2006; Petersen and Hughes 2007; Halling et al. 2008; Moncalvo and
Buchanan 2008). However, relatively little is known about the biogeography of Pluteus in its evolutionary
context other than conjecture in scattered reports such as those of Singer (1959) and Horak (1983). Progress
towards a global phylogeny of Pluteus has demonstrated the existence of a number of species complexes and
suggested surprising synonymies (Justo et al. 2011a; b). The reliability of numerous morphological characters
that have been used traditionally to distinguish species in Pluteus, such as colouration of the pileus has been
challenged by this work (Justo et al. 2011b). Some phylogenetic diversity appears to be correlated with patterns
of geographic distribution (Justo et al. 2011a; b), but the significance of this correlation, especially in regard to
taxonomy, has not been fully explored.
In order to further address biogeographic patterns within Pluteus and species delimitation within this genus,
we initiated a study of three exemplar species having putative multicontinental and disjunct distributions, viz.
P. albostipitatus, P. glaucotinctus and P. longistriatus. DNA sequences of the ITS region from numerous
collections from many countries were obtained and sequence divergences were calculated. Morphological data
were also gathered from numerous collections of these species, including the type specimens of each species.
Based on a synthesis of these data, we present a discussion concerning the geographical distribution and its
correlation to morphology and molecular data for these exemplar Pluteus species.
Capítulo II - 51
Material and methods
Taxon sampling and morphological study
The studied collections include specimens of P. albostipitatus from Africa, North and South America;
P. glaucotinctus from Africa, Asia, Europe and South America; and P. longistriatus from North and South
America (Fig. 1). The morphological studies were done using a total of 67 collections (35 of P. albostipitatus, 11
of P. glaucotinctus and 21 of P. longistriatus), which included the holotype of each species.
Fig. 1 Global distribution of P. albostipitatus, P. glaucotinctus and P. longistriatus. Filled stars indicate collections examined in this study;
empty stars indicate bibliographical records (see references in text); stars with black edges indicate the type locality for each species. The
types of the heterotypic synonyms of P. albostipitatus are indicated as 1. holotype of P. albostipitatus var. poliobasis; 2. holotype of
P. phaeoleucus; 3. holotype of P. melanopotamicus; 4. holotype of P. densifibrillosus. Collections with overlapping locations are not
indicated with multiple stars.
Colour terms follow Küppers (1979) and Munsell Color (2009). For microscopic analyses, the dried material
was wetted with 70% ethanol and then rehydrated in 5% KOH or stained with Melzer’s reagent. The notation
[a/b/c] at the beginning of a set of basidiospore data is to be read as “a basidiospores were measured from b
basidiomata taken from c collections”. Q represents the range of the length/width quotient for all the measured
spores, Qm represents the average of all computed Q values for all the measured basidiospores and Lm (Wm)
represents the average of all lengths (widths) of the measured basidiospores. At least 20 basidiospores from each
basidioma were measured in lateral view, and the terms denoting basidiospore shape follow Bas (1969). Generic
and infrageneric concepts follow Singer (1986) as complemented by Justo et al. (2011a; b). Herbarium acronyms
follow Thiers (2013).
Our evaluation of morphological, ecological, and geographical variations is also based, in part, on many
previous studies on these species (Peck 1878; 1885; Longyear 1904; White 1910; Kauffman 1915; 1918; Murrill
1917; Morse 1918; Sumstine 1918; Dennis 1953; Singer 1956; 1959; Rick 1961; Groves and Macrae 1963;
Horak 1964; Singer 1973; Gilbertson et al. 1974; Horak 1977; Horak and Heinemann 1978; Reid et al. 1981;
Pegler 1983; Branson 1989; Singer 1989; Courtecuisse 1991; Stijve and de Meijer 1993; Pegler 1997; Rodríguez
Capítulo II - 52
and Guzmán-Dávalos 2000; 2001; Meijer 2001; 2006; Wartchow et al. 2006; Bates 2006; Menolli and Capelari
2010; Menolli et al. 2010; Justo et al. 2011a; b; Gugliotta et al. 2012).
Molecular methods and phylogenetic analyses
The molecular analyses were completed with existing DNA sequences used by Justo et al (2011a; b) and from
newly collected sequences of materials representing at least one collection of each species from each
geographical area (Table 1). Methods for DNA isolation, PCR and sequencing follow Justo et al. (2011a; b). The
ITS region was amplified using the primer pair ITS1-F and ITS4 (White et al. 1990; Gardes and Bruns 1993).
Newly obtained sequences were deposited in GenBank. Molecular analyses were conducted using sequences
from sect. Pluteus, including those from the salicinus/albostipitatus clade, and the tree was rooted using
sequences of the cervinus/petasatus clade as outgroup (Table 1; clade names follow Justo et al. 2011a).
Sequences were aligned using MAFFT version 6 (http://mafft.cbrc.jp/alignment/server/; Katoh and Toh 2008)
with the Q-INS-I option. Then, the alignment was visually examined and manually corrected using MacClade
4.05 (Maddison and Maddison 2002). The alignment has been deposited in TreeBASE (S13390). Sequence
divergence was calculated using MatGAT (Campanella et al. 2003).
Table 1: Collection data and GenBank accession number of the Pluteus taxa used in the molecular analyses. Synonyms are indicated in
parentheses and represent the names as deposited in GenBank. The symbol "!" indicates a type collection.
Species
P. albostipitatus
P. albostipitatus
P. albostipitatus
P. albostipitatus
P. albostipitatus
P. albostipitatus
P. albostipitatus (as P. melanopotamicus)
P. albostipitatus (as P. densifibrillosus)
P. albostipitatus (P. phaeoleucus)
P. albostipitatus
P. albostipitatus
P. albostipitatus
P. cervinus (Schaeff.) P. Kumm.
P. glaucotinctus
P. glaucotinctus
P. glaucotinctus
P. glaucotinctus
P. glaucotinctus
P. glaucotinctus
P. glaucotinctus
P. harrisii Murrill
P. harrisii
P. harrisii (as P. puttemansii)
P. petasatus (Fr.) Gillet
P. salicinus (Pers. : Fr.) P. Kumm
P. salicinus
P. salicinus
P. salicinus
P. salicinus
P. saupei Justo & Minnis
ITS
FJ816656
FJ816661
JQ065032
JQ065033
JQ801375
JQ801373
JQ065031
HM562159
HM562141
HM562130
HM562106
HM562057
HM562035
HM562131
HM562132
HM562147
HM562157
JQ801374
JQ065023
JQ065024
FJ816652
FJ816654
HM562164
HM562038
JF908625
HM562051
JN603199
FJ774087
HM562174
HM562113
Collection (Herbarium)
NMJ128 (SP)
FK782 (SP)
FK1973 (SP)
FK1664 (SP)
FK1799 (SP)
FK1891 (SP)
R. Singer B12426! (F)
NMJ129! (SP)
GF5102! (BR)
GF5374 (BR)
AJ187 (MICH)
AJ154 (MICH)
AJ82 (LOU)
GF5274! (BR)
Thoen5546 (BR)
LASR64 (SP)
MC4412 (SP)
Pradeep13844 (CUW)
ARAN-Fungi 300824-1
ARAN-Fungi 300824-2
FK1066 (SP)
NMJ122 (SP)
NMJ131! (SP)
AJ201 (LOU)
MA67874 (MA)
AJ349 (CUW)
LE215427 (LE)
SF2 (BPI)
SGSaupe 230! (ILLS)
Country
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
Brazil
D. R. Congo
D. R. Congo
USA
USA
Spain
D. R. Congo
D. R. Congo
Brazil
Brazil
India
Spain
Spain
Brazil
Brazil
Brazil
Spain
Italy
Spain
Spain
Russia
USA
USA
Reference
Menolli et al. (2010)
This paper
This paper
This paper
This paper
This paper
Justo et al. (2011a; b)
This paper
This paper
This paper
--Justo et al. (2011a; b)
--Malysheva et al. (2009)
Justo et al. (2011a, b)
Justo et al. (2011a, b)
Capítulo II - 53
Maximum Parsimony (MP), Maximum Likelihood (ML) and Bayesian Analysis (BA) were performed with
the parameters specified in Justo et al. (2011b). The following abbreviations are used: Most Parsimonious Trees
(MPT), Bootstrap (BS) and Posterior Probability (PP).
Results and Discussion
Phylogenetic analyses and sequences divergence
The final data set consisted of 626 characters (gaps included) of which 158 are parsimony informative. In the MP
analysis, 6 MPT were recovered (Length = 332; CI = 0.71; RI = 0.91). One of the 6 MPT is shown in Fig. 2 and
it supports the discussions for P. albostipitatus and P. glaucotinctus. For the ITS topology of P. longistriatus see
Fig. 3 in Justo et al. (2011b).
3
JQ065033 Pluteus albostipitatus coll. FK1664 Brazil
Brazil
1
1
FJ816661 Pluteus albostipitatus coll. FK782 Brazil
Brazi l
Congoo
1 HM562141 Pluteus phaeoleucus coll. G-F 5102 (!) D.R. Cong
HM562130 Pluteus albostipitatus coll. G-F 5374 D.R. Congo
1
Pluteus albostipitatus
11
1
1
Brazi l
JQ065031 Pluteus melanopotamicus
melanopotamicu s coll. Singer B12426 (!)
(!) Brazil
Brazil
1
22
2
HM562159 Pluteus densifibrillosus
densifibrillosus coll. NMJ129 (!) Brazil
Brazil
FJ816656 Pluteus albostipitatus coll. NMJ128 Brazil
Brazil
100/100/1
6
HM562057 Pluteus albostipitatus coll. AJ154 USA
USA (FL)
(FL)
4
94/86/-
18
2
HM562106 Pluteus albostipitatus coll. AJ187 USA (FL)
(FL)
-/95/0.91
18
20
100/100/1
80/95/0.92
2
3
1
HM562164 Pluteus harrisii Brazil
HM562113 Pluteus saupei USA (IL)
HM562051 Pluteus salicinus Spain
HM562174 Pluteus salicinus USA (MI)
JN603199 Pluteus salicinus Spain
31
20
FJ816666 Pluteus harrisii Brazil
1
21
16
1 FJ816654 Pluteus harrisii Brazil
3
100/100/1
1
32
JF908625 Pluteus salicinus Italy
FJ774087 Pluteus salicinus Russia
100/100/1
Spain
pain
3 JQ065023 Pluteus glaucotinctus coll. ARAN 300824-1 S
JQ065024 Pluteus glaucotinctus coll. ARAN 300824-2 S
Spain
pain
6
HM562147 Pluteus glaucotinctus coll. LASR64 Brazil
79 1
HM562157 Pluteus glaucotinctus coll. MC4412 Brazi
Brazill
/83/5
JQ801374
Pluteus
glaucotinctus
coll.
Pradeep
13844 India
84/77/10
36
100/100/1
19
23
HM562035 Pluteus cervinus
HM562038 Pluteus petasatus
3
10
HM562132 Pluteus glaucotinctus coll. Thoen 5546 D.R. Con
Congo
go
HM562131 Pluteus glaucotinctus coll. Goossens-Fontana 5274 (!)
(!) D.R.
D.R. Congo
Congo
Pluteus glaucotinctus
Fig. 2 One of the 6 MPT from the phylogenetic analysis of selected Pluteus sect. Pluteus species. Branch lengths are shown above the
branches and support values below: BS from MP / BS from ML / PP from BY. Only BS > 70 and PP > 0.90 are presented. The symbol "!"
indicates a type collection.
Based on the current sampling, specimens of P. albostipitatus segregated into two clades, one containing
specimens collected in the USA and the second composed of collections from Brazil and Democratic Republic
of Congo (Fig. 2). No clear phylogeographic patterns were observed between collections from Brazil and
Capítulo II - 54
Democratic Republic of Congo in the ITS topology. For P. glaucotinctus, the sequence from the holotype
(Goossens-Fontana 5274) was recovered as sister to the remaining P. glaucotinctus sequences (Fig. 2). The
clade with collections from Spain, India and Brazil does not have geographically supported subclades and the
sequence from the other African specimen (Thoen 5546) fell on a branch external to the clade of Spain, India and
Brazil (Fig. 2). For P. longistriatus (> 90% BS in both MP and ML analyses) the sect. Hispidoderma tree
presented by Justo et al. (2011b) showed the specimens separated into two well-supported clades, the USA
branch and the clade composed of collections from Brazil (> 70% BS in both analyses).
In agreement with the phylogenetic analyses, the values of sequence divergence for P. albostipitatus (Table
2) showed that the higher values of sequence divergence occur between collections from the USA and all others,
varying between 1.80% and 3.48%. The sequences from the other localities differ themselves, in general, by
sequence divergence values between 0.30% and 1.34%, including almost identical sequences from distant
geographical areas (e.g. FK1973 from Brazil and GF5374 from Democratic Republic of Congo are 99.70%
identical). At times, collections from the same locality have lower similarity than collections from different
continents, e.g. FK782 and NMJ129 (holotype of P. densifibrillosus) are 98.67% identical. For P. glaucotinctus
(Table 3) the higher values of sequence divergence (varying from 4.18% to 4.95%) occur between the holotype
and all other collections. The other sequences in general have less than 2.43% sequence divergence and some
almost identical sequences were verified to occur in distant geographic areas (e.g. LASR64 from Brazil is 99.56%
and 99.09% identical to the Spanish and Indian collections, respectively). Finally, the sequence similarity
between the P. longistriatus collections is very high, with only 0.60% divergence between the USA and the three
Brazilian collections, which are identical to each other.
AJ187 (USA)
GF5374 (D. R. Congo)
NMJ129 (Brazil)
NMJ128 (Brazil)
FK782 (Brazil)
Singer B12426 (Brazil)
FK1664 (Brazil)
FK1891 (Brazil)
FK1799 (Brazil)
AJ187 (USA)
NMJ129 (Brazil)
NMJ128 (Brazil)
FK782 (Brazil)
Singer B12426 (Brazil)
FK1664 (Brazil)
FK1891 (Brazil)
FK1799 (Brazil)
FK1973 (Brazil)
AJ154 (USA)
Table 2: Sequence divergence (%) between all P. albostipitatus collections used in the molecular and phylogenetic analyses.
2.24
3.00
2.89
3.00
2.34
3.29
2.77
3.48
2.70
2.79
3.00
2.37
2.41
2.37
1.80
2.92
2.28
2.85
2.19
2.26
2.37
0.64
0.89
0.36
0.67
0.49
0.74
0.51
0.52
0.30
1.29
0.36
0.69
0.65
1.13
0.67
0.52
0.64
0.36
1.33
0.82
1.34
1.01
1.05
0.89
0.67
0.36
1.08
0.54
0.36
0.54
0.52
0.83
1.07
1.07
0.67
0.98
0.51
0.52
0.49
1.18
1.22
0.73
0.53
0.68
0.70
Capítulo II - 55
Thoen5546 (D. R. Congo)
LASR64 (Brazil)
MC4412 (Brazil)
ARAN300824_1 (Spain)
Thoen5546 (D. R. Congo)
LASR64 (Brazil)
MC4412 (Brazil)
Pradeep13844 (India)
Table 3: Sequence divergence (%) between all P. glaucotinctus collections used in the molecular and phylogenetic analyses.
4.18
4.62
4.76
4.22
4.22
4.95
1.76
1.76
2.06
2.06
2.43
0.29
0.44
0.44
0.91
0.74
0.74
1.21
0.00
1.37
1.37
Taxonomy and biogeographic implications
Pluteus albostipitatus (Dennis) Singer, Lloydia 21: 240. 1959.
(Figs. 3–5)
≡ Pluteus spilopus var. albostipitatus Dennis, Bull. Soc. Mycol. Fr. 69: 195. 1953.
= Pluteus albostipitatus var. poliobasis Singer, Beih. Sydowia 7: 64. 1973.
= Pluteus phaeoleucus E. Horak, Bull. Jard. Bot. Belg. 47: 89. 1977.
= Pluteus melanopotamicus Singer, Fieldiana, Bot. 21: 96. 1989.
= Pluteus densifibrillosus Menolli & Capelari, Mycologia 102: 698. 2010.
Pileus 10–60 mm diam, plane-convex or conic-campanulate, sometimes deeply concave, slightly to deeply
umbonate and slightly depressed at centre or around the umbo, light brown to greyish brown (Küppers –
N60A50M40 to N80A40M50; Munsell Color – 7.5YR 5/8, 6/8 or 10YR 5/1, 5/2, 6/1, 6/2), darker with coffee shades
at the centre (Küppers – N90A50M60; Munsell Color – 7.5YR 4/6), not hygrophanous, surface varying from
densely fibrillose to completely smooth, in the first case covered by appressed and radially arranged fibrils, more
concentrated at the centre and appearing radially split outside the central disc, sometimes exposing the context
which becomes translucently pinkish between fibrils of the marginal zone due to lamellar colour, finely pruinose,
fibrillose-punctate or with granular punctation at centre, margin usually sulcate-striate up to one-third or half of
the pileus radius, sometimes neither sulcate nor striate. Lamellae free, white, then pinkish (Küppers –
N10A20M30), crowded, ventricose, up to 6 mm broad, with whitish or concolourous edges and lamellulae of
different lengths, usually one lamellula for each lamella. Stipe 14–90 × 2–5(apex)–3–8(base) mm, slightly
tapering towards the apex, central, occasionally flexuous, cylindrical, white to pale cream, sometimes with
greyish to slightly silver base or almost dark grey, slightly longitudinally striate or completely smooth, with
scanty basal mycelium. Odour, taste, and context colour not recorded.
Basidiospores [704/35/35] (5.0–)5.5–9.5(−11.2) × (4.3–)5.0–9.0(−10.0) μm [Q = 1.00–1.24(−1.40−1.61);
Qm = 1.13; Lm = 6.9 μm; Wm = 6.2 μm], globose to broadly ellipsoid, rarely ellipsoid or elongate, inamyloid,
hyaline, smooth, thick-walled, sometimes guttulate. Basidia 19.0–35 × 7.0–15.0 μm, versiform to clavate or
narrowly utriform, thin-walled, 4-spored, sometimes with small scattered guttules. Pleurocystidia 27–96 × 10.0–
32 μm, fusiform or slightly ventricose to slightly utriform, sometimes (slightly) clavate, with rounded,
Capítulo II - 56
Fig. 3 Basidiomata of P. albostipitatus. a. FK1891; b–c. FK1909; d. FK1598; e–f. FK1973; g. FK1061; h–i. NMJ129 (holotype of
P. densifibrillosus – from Menolli and Capelari 2010); j–k. NMJ128; l. FK1796; m. FK2147; n. AJ187; o–p. (from Horak and Heinemann
1978); o. Goossens-Fontana 245/888; p. Goossens-Fontana 5102 (holotype of P. phaeoleucus). All scale bars = 1 cm.
subcapitate or truncate apex, some with an apical constriction like a neck and forming two rounded and
versiform lateral projections, with an internal septum in the upper part (observed frequently in one North
American collection), and rarely with 2-4 well developed prongs at apex or with several small finger-like
Capítulo II - 57
protuberances (in a few Brazilian collections), thin- to slightly thick-walled, sometimes like metuloids with wall
thickened up to 2 μm wide at upper part or wall evenly thickened, colourless, hyaline, moderately numerous and
common all over lamellar faces. Cheilocystidia 25–96(–108) × 7.0–29 μm, mostly clavate with a well developed
pedicel at the base, some narrowly ovoid or rarely slightly ventricose to lageniform like pleurocystidia (the latter
type was observed only in few Brazilian collections), thin-walled, colourless, hyaline, abundant, scattered or
clustered and forming a well-developed strip. Lamellar edge sterile, completely covered with cystidia organized
in fascicules or heterogeneous with a mixture of cystidia and basidioles. Lamellar trama inverse, up to 62 μm
wide, composed of thin-walled hyphae, 2.5–12.5 μm diam, hyaline. Pileus context undifferentiated, composed of
thin-walled hyphae, 3.7–18.7 μm diam, hyaline. Pileipellis a cutis up to 110 μm thick, with few ascendant
subradial elements, composed of thin or slightly thick-walled hyphae, individual terminal elements 65–120 ×
10–20 μm, cylindrical or fusiform, with rounded to subacute apex (up to 5.0 μm diam), mostly with dissolved
brown intracellular pigment. Stipitipellis a cutis of thin-walled hyphae, 5.0–12 µm wide, colourless and mostly
hyaline. Clamp connections absent in all parts examined.
Type description (holotype of P. spilopus var. albostipitatus) Basidiospores [20/1/1] (6.0–)7.5–8.7(–10.0) ×
6.2–7.5(–8.7) µm (Q = 1.00–1.21 Qm = 1.08; Lm = 7.8; Wm = 7.3), globose, subglobose or broadly ellipsoid,
inamyloid, hyaline, smooth, slightly thick-walled, usually one-guttulate. Basidia not observed. Pleurocystidia
37–75 × 12.5–21 µm, fusiform or slightly ventricose to slightly utriform, sometimes slightly clavate, with
rounded, subcapitate or truncate apex, slightly thick-walled up to 1 μm wide, colourless, hyaline, moderately
numerous and common all over lamellar faces. Cheilocystidia 41–72 × 10.0–17.5 μm, clavate with a well
developed pedicel at the base, thin-walled, colourless, hyaline, abundant, clustered forming a well-developed
strip. Lamellar edge sterile, completely covered with cystidia organized in fascicules. Pileipellis a cutis but
difficult to describe the hyphal elements. Clamp connections absent in all parts examined.
Material examined BRAZIL: Rick s.n. (BPI770882 as P. murinus Bres.); Amazonas State, Igarapé do
Tarumãzinho, 31 March 1983, Singer B12426 (F – holotype of P. melanopotamicus); Bahia State, Itabuna city,
12 June 1978, Singer B11231 (INPA); Minas Gerais State, Alto Caparaó city, Parque Nacional do Caparaó, 18
March 2011, F. Karstedt FK1973 (SP); Nova Ponte city, Fazenda Caxuanã, 22 January 2007, R. Maziero s.n.
(SP417354); R. Maziero s.n. (SP417355); R. Maziero s.n. (SP417356); R. Maziero s.n. (SP417357); R. Maziero
s.n. (SP417358); R. Maziero s.n. (SP417359); Pará State, Belém city, Ilha do Combu, 20 January 2011,
F. Karstedt FK1891 (SP); Paraná State, Antonina city, Reserva Natural do Rio Cachoeira, 24 May 2003, A.A.R.
de Meijer ANc-4219 (MBM); Foz do Iguaçu city, Parque Nacional do Iguaçu, 7 November 2010, F. Karstedt
FK1664 (SP); Mandirituba city, 20 April 1993, A.A.R. de Meijer MANa-2629 (MBM); Paranaguá city, Mundo
Novo, 27 May 1993, A.A.R. de Meijer PAc-2744 (MBM); Rio Grande do Sul State, Guaíba city, Horto Florestal
da Conpanhia Estadual de Energia Elétrica, 3 October 1984, J. Medeiros s.n. (ICN102101); Passo Fundo city,
Universidade de Passo Fundo, 30 October 2003, M.S. Rother & B.M.A. Severo s.n. (RSPF284); São Francisco de
Paula city, Floresta Nacional do Instituto Brasileiro de Desenvolvimento Florestal, 19 March 1980, A. Batista
s.n. (ICN6834); São Salvador city, 1944, Rick s.n. (PACA20770 as P. sensitivus var. macrospora nom. in herb.);
Santa Catarina State, Gaspar city, Reserva Particular do Patrimônio Natural Figueira Branca, 30 July 2010,
F. Karstedt FK1598 (SP); 6 January 2012, F. Karstedt FK2147 (SP), Blumenau city, Parque Nacional Serra do
Itajaí, Parque das Nascentes, Trilha da 3ª Vargem, 25 January 2011, F. Karstedt & R. Karstedt FK1796 (SP);
Capítulo II - 58
Fig. 4 Pluteus albostipitatus. a–f. Basidiospores; g–r. Pleurocystidia; a, g. FK1664; b, k. FK1973; c, m. Singer B12426 (holotype of
P. melanopotamicus); d, n. FK1891; e, o. NMJ129 (holotype of P. densifibrillosus); f, p. NMJ128; h. FK782; i. Goossens-Fontana 5102
(holotype of P. phaeoleucus); j. Goossens-Fontana 5374; l. FK1799; q. AJ154; r. AJ187. All scale bars = 10 µm.
F. Karstedt & R. Karstedt FK1799 (SP); São Paulo State, Cananéia city, Parque Estadual da Ilha do Cardoso,
restinga, 17 November 1982, G. Guzman et al. 22947 [SP as P. cubensis (Murrill) Dennis]; Mogi Guaçu city, 30
January 1987, D.N. Pegler 3890 [K(M) as P. sergii Singer]; São Paulo city, Parque Estadual da Cantareira,
Capítulo II - 59
Núcleo Engordador, 23 October 2006, F. Karstedt & M. Capelari FK782 (SP); 24 April 2007, Menolli Jr. et al.
NMJ129 (SP – holotype of P. densifibrillosus); 24 July 2007, Menolli Jr. et al. NMJ128 (SP); 31 January 2008,
F. Karstedt et al. FK1061 (SP); Parque Estadual das Fontes do Ipiranga, 1 March 2011, F. Karstedt & J.J.S.
Oliveira FK1909 (SP). DEMOCRATIC REPUBLIC OF CONGO: Panzi-Kivu, 1650 m altitude, March 1954,
Goossens-Fontana 5374 (BR); December 1949, Goossens-Fontana 5102 (BR – holotype of P. phaeoleucus).
TRINIDAD: Diego Martin, 29 September 1949, R.W.G. Dennis 89 [K(M) – holotype of P. spilopus var.
albostipitatus]. USA: Florida, Tallahassee, Lake Overstreet, 24 August 2009, A. Justo AJ154 (MICH); 28 August
2009, A. Justo AJ187 (MICH).
Habit and habitat Solitary, in small groups or subgregarious, on decayed branches and trunks of dicotyledons
or on decayed stubs of palm tree.
Geographical distribution Pluteus albostipitatus was described from Trinidad (Dennis 1953 as P. spilopus
var. albostipitatus) and it is known for several countries in the Americas and Africa (Fig. 1): Argentina (Singer
1956; 1959; Horak 1964), Bolivia (Singer 1959), Brazil (Stijve and de Meijer 1993 as P. cf. albostipitatus;
Meijer 2001; 2006; Wartchow et al. 2006; Pegler 1997; Menolli et al. 2010), Democratic Republic of Congo
(Horak and Heinemann 1978), Ecuador – Galapagos Islands (Reid et al. 1981), French Guyana (Courtecuisse
1991), Martinique (Pegler 1983) and the USA (Justo et al. 2011a; b). Singer (1973) also described from Mexico
(Veracruz State) P. albostipitatus var. poliobasis, which was later recollected by Rodríguez and GuzmánDávalos (2000, 2001) in Jalisco State, Mexico. Pluteus albostipitatus is a widely distributed species and this
study expands the known geographical distribution of this species, especially in Brazil where it is known with
certainty from the states of Amazonas, Bahia, Minas Gerais, Pará, Paraná, Rio Grande do Sul, Santa Catarina
and São Paulo.
Fig. 5 Holotype of P. albostipitatus (R.W.G. Dennis 89). a. Basidiospores; b. Pleurocystidia; c. Cheilocystidia. All scale bars = 10 µm.
Discussion Pluteus albostipitatus is a species with considerable morphological plasticity without any
correlation between molecular and/or geographical distribution, and this plasticity led to this species being
described a number of different times. Its occurrence is restricted to subtropical and tropical areas where
basidiomata may develop year round. In countries where it is less well known, the fructification period seems to
be more restricted (e.g. February and March in Argentina; March in Bolivia and French Guyana; March and
Capítulo II - 60
December in Democratic Republic of Congo; June in Galapagos Islands; August in the USA; September in
Martinique and Trinidad). In Brazil, where the species is well known, fruiting has been documented in almost
every month (hitherto now except in February, August, September and December) and there does not appear to
be a distribution pattern that correlates with latitudinal position and months of the year. An increase in collecting
in the other countries will be necessary to better understanding the occurrence of fruiting, but we expect
P. albostipitatus produces basidiomata throughout the year in regions characterised by subtropical and tropical
climates.
Pluteus albostipitatus was classified originally in sect. Hispidoderma (Singer 1959), which is characterised
mainly by the presence of thin-walled pleurocystidia and a non-hymeniderm or non-epithelial pileipellis.
Although some published descriptions refer the pleurocystidia as non-metuloid and thin-walled, previous studies
(Singer 1959; Pegler 1983) pointed out a probable relationship between P. albostipitatus and members of sect.
Pluteus, a section that is characterised mainly by metuloid and horned pleurocystidia, since P. albostipitatus
possesses unusual pleurocystidial shape with the presence of small apical outgrowths. Recent morphological
studies (Wartchow et al. 2006; Menolli et al. 2010) reported the presence of pleurocystidia with slightly
thickened to thickened walls and some lateral or apical prongs in collections from Brazil. The position of
P. albostipitatus in sect. Pluteus is confirmed based on the present and previous molecular studies of Pluteus
phylogeny (Menolli et al. 2010; Justo et al. 2011a; b). It appears that the pleurocystidia represent a transitional
form within sect. Pluteus. Though they have a different origin than the pleurocystidia found in sect.
Hispidoderma, they are morphologically similar to those of sect. Hispidoderma and often include forms that are
intermediate in appearance between this type and those more typical of sect. Pluteus. These transitional forms
and variations can be easily observed by studying many collections as was done in the present study, where it
was observed that the shape of pleurocystidia varies from thin-walled with a rounded apex (Fig. 4o) to having
slightly thickened to thickened walls with some short lateral prongs (Fig. 4h, p, q) to having slightly thickened to
thickened walls with well-developed prongs (Fig. 4g, m).
The present morphological and molecular analyses allowed the synonymisation of P. albostipitatus var.
poliobasis described from Mexico (Singer 1973), P. phaeoleucus from Democratic Republic of Congo (Horak
1977; Horak and Heinemann 1978), P. melanopotamicus (Singer 1989) and P. densifibrillosus (Menolli and
Capelari 2010; Justo et al. 2011b) both from Brazil. Based on our morphological studies, the collections reported
as P. aff. harrisii by Meijer (2006) and as P. cubensis (Guzmán et al. 22947) by Pegler (1997), as well as the
other one named as P. sensitivus var. macrospora (nom. in herb.) and reported as P. sensitivus Rick (Rick 1961)
are actually P. albostipitatus.
The several taxa that are now considered synonyms of P. albostipitatus were proposed as new on the basis of
the stipe colour variation, the pileus shape, the morphological variation of pleurocystidia and the shape and size
of basidiospores. For instance, P. albostipitatus var. poliobasis differs from the type variety only by the pearlgray colour (griseo) of the stipe (Singer 1973). However, colour variation tends to display frequent
morphological plasticity within some species of Pluteus and since this character has a wide range of variation
and intergrades to a great extent between many fresh collections that have been observed, P. albostipitatus var.
poliobasis is considered herein a synonym of P. albostipitatus. Pluteus phaeoleucus was described as a new
species based on the absence of pleurocystidia (Horak 1977; Horak and Heinemann 1978), which were later
observed by Justo et al. (2011b) and illustrated herein (Fig. 4i). Pluteus melanopotamicus was described as a
new species mainly due to the occurrence of pleurocystidia with several rounded and small finger-like
protuberances at the apex and the presence of dimorphic basidiospores, which differ in size by a measurement
Capítulo II - 61
interval of just 0.5 µm in length and 1.0 µm in width between the two types of basidiospores (Singer 1989).
Finally, P. densifibrillosus was described as distinct from P. albostipitatus by deeply concave pileus and by the
almost globose basidiospores (Menolli and Capelari 2011), which in P. albostipitatus were commonly classified
with a wide variation in shape and size or even in three classes (small, normal and giant) but never globose
(Singer 1959).
Although the USA collections have a high level of molecular divergence when compared to the other
collections (1.80% and 3.48%), we do not feel there are enough molecular and morphological differences to
accept them as distinct species at this time. The only distinctive character observed in one North American
collection (AJ187) was the presence of some pleurocystidia with an internal septum in the upper part (Fig. 4r).
However, this character is probably not limited to populations in the USA. Singer (1959) also described
pleurocystidia as “exceptionally with a pseudo-(secondary) septum near the tip” in collections from Central and
South America.
As previously pointed out by Justo et al. (2011b), P. albostipitatus can be considered a morphologically
cryptic species since there is no correlation between the geographical distribution, morphology and molecular
variation. Additional collections from other localities in North America could confirm the existence of a second
phylogenetic species within the same morphological species concept. New collections of P. albostipitatus from
western, southern or southeastern Asia and Oceania would demonstrate a pantropical distribution for this
species.
Pluteus glaucotinctus E. Horak, Bull. Jard. Bot. Belg. 47: 88. 1977.
(Figs. 6, 7)
Pileus 20–90 mm diam, conic-campanulate at first, expanding to convex or plane, sometimes with a broad
obtuse umbo that becomes depressed at centre with age, surface at least one-half of the radius towards margin
varying from white to light beige or slightly pinkish, smooth (in some cases apparently smooth but slightly
fibrillose under lens) except at centre which has olivaceous (Munsell Color – 2.5Y 6/6, 6/8, 7/6, 7/8) to dark grey
or bluish grey (Munsell Color – in the range of Gley2 4/1, 5/1, 6/1) squamules and fibrils, becoming fibrillose to
densely fibrillose all over in older specimens, slightly hygrophanous, margin usually striate up to 1/3 of the
pileus radius, sometimes indifferently or deeply striate. Lamellae free, white then pinkish (Munsell Color – 7.5R
8/4), crowded, ventricose, up to 7 mm broad, with whitish and flocculose or concolourous edges and lamellulae
of different lengths, one lamellula for each 1–2 series of lamellae. Stipe 35–150 × 2–10 mm, cylindrical (or
compressed, but probably due to an irregular development in the substrate), subequal and slightly tapering
towards the apex, central, white to pale cream at apex with olive green fibrils, light brown to slightly greenish
punctations or distinct blue to blue-green tinges (the latter only recorded in Spanish and African collections) at
base, slightly longitudinally striate and usually with scanty basal mycelium. Context in stipe and pileus white, or
in some places with slight blue-green tinges (in Spanish collections). Odour indistinct and taste indistinct to
slightly sweet. Spore print brownish-pink (Munsell Color – 2.5YR 6/6, 6/8).
Basidiospores [254/11/11] 5.0–10.0 × 5.0–7.5 μm [Q = 1.10–1.61(−1.74); Qm = 1.32; Lm = 7.8 μm; Wm =
6.0 μm], subglobose to ellipsoid, rarely elongate, inamyloid, hyaline, smooth, thick-walled, usually guttulate.
Basidia 17.5–36 × 6.2–18.0 μm, versiform to clavate or narrowly utriform, thin-walled, 4-spored, sometimes
with small scattered guttules. Pleurocystidia (30–)40–85 × (7.0–)10.0–28 μm, fusiform to lageniform, subclavate
to clavate, or slightly ventricose to slightly utriform, usually with a rounded apex or with an almost truncate
apex, which can be slightly constricted or with minor apical and lateral projections (only in two Brazilian
Capítulo II - 62
collections and like the poorly developed prongs observed in P. albostipitatus), thin-walled, a few with some
incrusting parietal pigment at apex (in Indian collection), colourless, hyaline, scattered and relatively common all
over lamellar faces. Cheilocystidia 35–94 × 6.2–28 μm, slightly ventricose to lageniform, usually elongated to
slender with a well developed pedicel, sometimes clavate or mucronate, apex rounded or usually with a narrow
and short papilla, rarely truncated, thin-walled, a few with some incrusting parietal pigment at apex (in Indian
collection), colourless and hyaline or with pale brown intracellular pigment (observed in few cheilocystidia of
the holotype), abundant and usually forming a well-developed strip. Lamellar edge sterile, completely covered
with cystidia. Lamellar trama inverse, up to 50 μm wide, composed of thin-walled hyphae, 2.5–8.7 μm diam,
hyaline and rarely clamped (in Indian collection). Pileus context undifferentiated, approx. 360 μm thick,
composed of thin-walled hyphae, 3.7–11.0(–16.0) μm diam, colourless and hyaline, sometimes with blue-green
pigment (in Spanish collections) or with oleiferous hyphae. Pileipellis a cutis of approx. 90 μm thick, composed
of thin-walled hyphae, individual terminal elements 55–380 × 5.0–18.0(–23) μm, cylindrical or fusiform, with
rounded to subacute apex, colourless or with dissolved dirty pale to brown intracellular pigment, a blue-green
intracellular pigment and sometimes an additional parietal pigment were also seen in the Spanish collection.
Stipitipellis a cutis of cylindrical hyphae, 3.5–15.0 µm wide, thin-walled, colourless or with brown intracellular
pigment; in Spanish collections it is possible to see distinct terminal elements at apex (75–138 × 12.0–16.0 µm,
narrowly clavate or fusiform) and flexuous hyphae filled with blue-green or greyish pigment at base. Clamp
connections absent in all parts examined, except for those rarely seen in the thin-hyphae of the lamellar trama in
the Indian collection.
Type description Basidiospores [30/1/1] 6.5–9.0 × 5.0–6.5 μm, (Q = 1.15–1.55; Qm = 1.33; Lm = 7.6 μm; Wm =
5.7 μm), broadly ellipsoid to ellipsoid, inamyloid, hyaline, smooth, slightly thick-walled. Basidia not observed.
Pleurocystidia 50–70 × 15.0–22 μm, lageniform or narrowly utriform, thin-walled, colourless, hyaline, scattered
to relatively common all over lamellar faces. Lamellar edge sterile, completely covered with cystidia.
Cheilocystidia 45–61 × 12.0–17.0 μm, lageniform, with rounded apex or some with truncate apex, thin-walled,
colourless or (a few) with pale brown intracellular pigment, crowded and forming a well-developed strip.
Pileipellis a cutis composed of thin-walled hyphae, individual terminal elements 70–90 × 12.0–17.0 μm,
cylindrical or fusiform, colourless and hyaline or with brown intracellular pigment. Stipitipellis a cutis of thinwalled hyphae, 5.0–12.0 μm wide, colourless and hyaline. Clamp connections absent in all parts examined.
Material examined BRAZIL: São Paulo State, Santo André city, Reserva Biológica do Alto da Serra de
Paranapiacaba, 31 October 1990, L.K. Okino & A.M. Glugliotta 3630 (SP as P. atriavellaneus Murrill); São
Paulo city, Parque Estadual da Cantareira, Núcleo Engordador, 19 February 2008, F. Karstedt et al. FK1084
(SP); Parque Estadual das Fontes do Ipiranga, 2 December 2008, M. Capelari & U.C. Peixoto MC4412 (SP); 27
October 2009, L.A.S. Ramos LASR64 (SP). DEMOCRATIC REPUBLIC OF CONGO: Panzi-Kivu, 1650 m
altitude, March 1953, Goossens-Fontana 5274 (BR – holotype); Kipopo, November 1972, Thoen 5546 (BR).
INDIA: Kerala State, Trivandrum District, Enikkara, 1 November 2011, Pradeep 13844 (CUW). SPAIN:
Guipúzcoa, Donostia/San Sebastián, 18 August 2011, leg. P. Arrillaga s.n. (ARAN-Fungi 3008424-1);
1 September 2011, leg. P. Arrillaga s.n. (ARAN-Fungi 3008424-2); 3 September 2011, leg. P. Arrillaga s.n.
(ARAN-Fungi 3008424-3); 11 September 2011, leg. P. Arrillaga s.n. (ARAN-Fungi 3008424-4).
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Habit and habitat Solitary, in small groups or subgregarious, on dead wood, inside coffee and Eucalyptus
plantations (Africa); on dead unidentified angiosperm wood (India); on brown-rotted log of Quercus sp. (Spain);
and on standing tree with fructification of other polyporoid fungi (Brazil).
Fig. 6 P. glaucotinctus. a–g. Basidiomata (Scale bar = 1 cm); h. Pileipellis elements in Congo Red (Scale bar = 10 µm). a. LASR64;
b–c. MC4412; d. ARAN-Fungi 3008424-3; e. ARAN-Fungi 3008424-4; f, h. ARAN-Fungi 3008424-1; g. ARAN-Fungi 3008424-2.
Geographical distribution Pluteus glaucotinctus was described from Democratic Republic of Congo (Horak
1977; Horak and Heinemann 1978). Recently, it was recorded from Brazil (Justo et al. 2011a; b) and now the
known distribution is expanded to Spain and India (Fig 1).
Discussion Pluteus glaucotinctus is apparently a pantropical species-complex that has been found in
association with a great variation of habitats (dead wood, brown-rotted log and living tree) over its range.
Considerable morphological differences and a substantial level of sequence divergence, mainly between the
holotype and the remaining collections, were also observed.
Considering the pantropical distribution, its occurrence in Spain seems to be an exception from this pattern,
which could be explained tentatively by the climatological conditions during the fructification. Data from the
nearest climatological station (Oiartzun Observatory, located less than 10 km away from the collecting site)
indicated that between mid-August and mid-September 2011 the daily average temperatures were between 17
and 26 C and the relative humidity was between 70 and 97%, which are favorable conditions for tropical fungal
fructification.
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Regarding morphology, some distinctive characters were evident that might distinguish individual specimens
such as cheilocystidial colouration, variations in pleurocystidia, the presence of blue-green pigments, and clamp
connections. The presence of cheilocystidia with pale brown intracellular pigment is a distinctive character that
is only observed in the holotype (Democratic Republic of Congo), but this is probably without correlation to the
geographic distribution because those found in the collection Thoen5546 from the same area are colourless. The
pigmentation or presence of content in cystidia is often not a good morphological character to separate related
taxa because in the same collection it is sometimes possible to find pigmented or colourless cystidia. Within the
range of molecular variation used to delimit species, it is also possible to find both pigmentation possibilities in
different collections as observed in the pleurocystidia of North and South American collections of
P. longistriatus (see present discussion under this species) and in cheilocystidia of P. eludens E.F. Malysheva,
Minnis & Justo (Justo et al. 2011c) as well as P. phlebophorus (Ditmar) P. Kumm., which includes material with
pigmented cheilocystidia that may be applied under the species concept of P. luctuosus Boud. (Justo et al.
2011b).
The arrangement of pileipellis allied with the presence of minor apical and lateral projections (Fig. 7e) in the
thin-walled pleurocystidia (like those observed in P. albostipitatus) could be used as morphological characters to
correctly classify P. glaucotinctus in sect. Pluteus. Although these projections were observed in only two
(FK1084 and LASR64) of the three Brazilian collections, they probably can not be considered a variation
correlated to a geographical area. Similar characters were also observed in many collections of P. albostipitatus
from different geographical localities and these also were without any molecular correlation.
Although the blue-green pigments (Fig. 6e–h) have been observed by us only in Spanish specimens, they
were recorded previously for the African collections (Horak 1977; Horak and Heinemann 1978) and this
character is probably common in P. glaucotinctus. It certainly correlates to the presence of psilocybin, which has
been recorded from other taxa in this clade such as P. salicinus (Pers. : Fr.) P. Kumm. (Saupe 1981, Stijve and
Bonnard 1986). Notably, the Saupe material (Saupe 1981) actually represents a distinct species, P. saupei Justo
& Minnis (Justo et al. 2011b). Stijve and Bonnard (1986) mentioned the presence of psilocybin in P. nigroviridis
Babos and later Stijve and de Meijer (1993) recorded it for P. glaucus Singer. Pluteus nigrolineatus Murrill is
also frequently reported with blue-green pigments at stipe base (Singer 1962, 1969; Rodríguez and GuzmánDávalos 2001) and although it was traditionally classified in sect. Hispidoderma (Singer 1986), Menolli et al.
(2010) recently showed by molecular data that it is positioned in sect. Pluteus clade. It has been demonstrated
that all of these species are grouped together in the salicinus/albostipitatus clade (Menolli et al. 2010; Justo et al.
2011b), which could indicate that in sect. Pluteus this character is restricted to species in this clade. However, to
confirm this homology it will be necessary to investigate the phylogenetic position of P. nigroviridis and
P. glaucus as well as the presence of psilocybin in other species of this clade, including P. harrisii Murrill and
P. albostipitatus. Furthermore, Pluteus species with blue-green tinges are also present in other sections such as
P. cyanopus Quél (Homola 1972) and P. phaeocyanopus Minnis & Sundb. (Minnis and Sundberg 2010) from
sect. Celluloderma, which indicates that this character has originated at least twice during the evolution of the
genus.
The Indian collection has a few noteworthy morphological differences including the presence of some
incrusting parietal pigment at the apex of a few pleuro- and cheilocystidia and clamp connections that were
rarely seen in the thin-hyphae of the lamellar trama. The presence of clamp connections is the most distinctive
character between Indian and other collections, and this might be significant given the use of this morphological
character to support the distinction of different complexes or clades of Pluteus (Justo et al. 2011b). However,
Capítulo II - 65
considering that the clamp connections were rare and barely observable, it is possible that they are present but
overlooked in the other collections. Clamp connections are often more difficult to find in older, dried herbarium
material.
Fig. 7 Pluteus glaucotinctus. a–c. Basidiospores; d–i. Pleurocystidia; j–o. Cheilocystidia; p. Pileipellis elements. a, d, j, p. ARAN-Fungi
3008424-1; b, e, k. LASR64; c, f, l. MC4412; g, m. Pradeep 13844; h, n. Thoen 5546; i, o. Goossens-Fontana 5274 (holotype). All scale bars
= 10 µm.
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The considerable habitat variation over the widespread distribution of P. glaucotinctus and the ecological
implications of this is worthy of comment. The fructification of the Spanish collections on a brown-rotted log is
particularly noteworthy. This habitat is anomalous for Pluteus, which is usually collected from white-rotted
wood or more rarely on other organic debris. Wood that has been decomposed by brown-rot decay is deprived of
most of its cellulosic components while the lignin remains almost intact, thus providing very little nutrients for
fungal species (Yelle et al. 2008; Martinez et al. 2009). While it is very unlikely that P. glaucotinctus was
responsible for the brown-rot decay observed in the substrate, it is highly remarkable that this fungus is not only
able to survive in such a nutrient-poor environment but that it is also able to produce basidiomata many times in
less than one month. No other fungal basidiomata were observed in the same log where P. glaucotinctus was
growing, but that does not exclude the possibility that other fungi were present at the same time or before the
arrival of P. glaucotinctus. In South America, one of the three Brazilian specimens (LASR64) was collected from
a standing tree (apparently a living tree) where a polyporoid fungus was also present and likely responsible for
the death of the plant tissue where P. glaucotinctus was growing. Despite these records, more collections of
Pluteus with similar habitats and habits are needed to establish a well-grounded discussion about the ecological
pattern within this species and the genus as a whole.
Considering all these individual differences among each collection, it is difficult to establish any
distinguishable features to reliably separate them or recognize any distinct taxa, especially when all
morphological, molecular and geographic variations are considered. Molecular data seem to indicate the
existence of three different species within the broad morphological concept outlined above: (i) Pluteus
glaucotinctus sensu stricto, represented solely by the holotype collection; (ii) a second putative species
represented by the collection Thoen5546 and (iii) a third species present in Brazil, India and Spain. Given the
extensive, and overlapping, morphological variation and considering the fact that two of the putative species in
this group are only represented by a single herbarium collections more than 50 years old we refrain from
proposing a formal subdivision of the P. glaucotinctus complex. Additional collections from Africa are needed
to resolve this species-complex.
Pluteus longistriatus (Peck) Peck, Ann. Rep. N.Y. St. Mus. Nat. Hist. 38: 137. 1885.
(Figs. 8–11)
≡ Agaricus longistriatus Peck, Ann. Rep. N.Y. St. Mus. Nat. Hist. 30: 40. 1878.
Pileus 12–42 mm diam, conic-convex or campanulate to plane-convex, ovoid when young (approx. 6-9 mm
diam), slightly umbonate and sometimes slightly depressed around the umbo, completely dark brown when
young (Küppers – A99M70C70), then pale fuscous to light brown-grey (Küppers – A80M60C60 to A99M60C60), centre
distinctly fibrillose, sometimes forming granules or floccons, but often appearing completely naked and
glabrous, dark brown fibrils more concentrated at the centre and more spaced and discolouring radially towards
the margin, margin deeply sulcate-striate over at least one half of the radius. Lamellae free, pinkish, crowded,
with lamellulae. Stipe 30–70 × 1–3(apex)–2–5(base) mm, central, hollow, slightly tapering towards the apex,
usually longitudinally striate, white to white-cream, sometimes with scanty brownish fibrils at base and white
basal mycelium. Context white or greyish-white at first with slightly more cinereous shades at base and near the
margin in young specimens, later usually white or pallid, moderately thin and fragile, unchanging when
wounded. Odour slightly acid-fungine or inodorous.
Basidiospores [372/21/21] 6.0–8.7 × (4.3–)5.0–7.5 µm (Q = 1.11–1.50; Qm = 1.24; Lm = 7.1 µm; Wm = 5.7
µm), subglobose, broadly ellipsoid or ellipsoid, inamyloid, hyaline or subhyaline to pale yellow in KOH,
Capítulo II - 67
smooth, thick- or slightly thick-walled, sometimes guttulate with one to several scattered guttules. Basidia 15–26
× 6.0–11.2 µm, versiform or clavate to fusoid-ventricose, thin-walled, 4-spored, contents granular or agranular.
Pleurocystidia 35–106(–114) × (8.7–)10–30(–38) µm, clavate or ventricose to lanceolate, with a rounded or
occasionally a narrow papillate apex, thin-walled, subhyaline with pale straw dissolved content or hyaline and
colourless without contents (majority of North American collections), numerous. Cheilocystidia 16–55(–61) ×
6.5–18(–22) µm, the majority similar in shape to pleurocystidia with some smaller or versiform-clavate,
pyriform to spheropedunculate, thin-walled, colourless, hyaline, moderately numerous to crowded. Lamellar
edge sterile. Lamellar trama inverse, up to 40 µm wide, composed of thin-walled hyphae, 2.5–6.2 µm diam, some
branched, colourless, hyaline, with oleiferous hyphae up to 5.0 µm diam. Pileus context undifferentiated,
approximately 69 mm thick, composed predominantly of oleiferous hyphae up to 7.5 µm diam, moderately
branched, thin-walled. Pileipellis a hymeniderm composed of one layer of versiform elements, terminal cells 35
–150(–170) × 8.0–43(–52) µm, vesiculose, clavate to fusoid cells [approx. 35–108(–117) × 8.0–43(–52) µm]
intermixed by elongate-fusiform or elongate-clavate to filiform cystidioid elements [approx. 105–150(–170) ×
8.0–33(–40) µm], apices typically rounded or rarely with a short papilla or mucro, thin-walled and with
dissolved or sometimes condensed brown intracellular or vacuolar contents. Stipitipellis a cutis of cylindrical
hyphae, 3.0–24 µm wide, thin-walled, hyaline, lacking apparent contents. Caulocystidia 32–120(–134) × 8.0–34
µm, clavate to elongate-clavate or fusoid, apices typically rounded or rarely with narrow and short papilla, thinwalled, hyaline, often with brown intracellular contents, scattered in fasciculate bundles, numerous or sometimes
absent in some North American collections. Clamp connections absent in all parts examined.
Fig. 8 Basidiomata of P. longistriatus. a–b. FK1660; c. FK718; d–e. LASR25 (d. from Menolli and Capelari 2010); f. Minnis 03-09-20-3. All
scale bars = 1 cm.
Type description Basidiospores [30/1/1] 6.0–7.5 × 4.5–7.0 µm (Q = 1.09–1.38; Qm = 1.25; Lm = 6.7; Wm =
5.4), predominantly broadly ellipsoid to less commonly subglobose or ellipsoid, inamyloid, subhyaline to pale
yellow in KOH, smooth, slightly thick-walled, contents typically absent probably due to age of the herbarium
material. Basidia 18.0–26 × 6.5–9.5 µm, clavate to fusoid-ventricose, thin-walled, 4-spored, contents typically
absent probably due to age of the herbarium material. Pleurocystidia 40–80 × 13.0–24 µm, cylindrical-clavate to
ventricose, with a rounded apex, thin-walled, hyaline and lacking apparent contents, scattered and numerous.
Cheilocystidia 16–35 × 6.5–13 µm, versiform-clavate, pyriform to spheropedunculate or ventricose to slightly
Capítulo II - 68
Fig. 9 Pluteus longistriatus. a–c. Basidiospores; d–m. Pleurocystidia; n–s. Cheilocystidia; t–z . Caulocystidia. a, e, o. FK681; b, i, r, x.
Minnis 03-09-20-3; c, k, y. M. Morris s.n.; d, n, t. FK680. f, p, u. LASR25; g, v. MC4431; h, q, w. H.E.B. 15915; j. D.R. Sumstine s.n., July
1921; l, s. T.J. Baroni & R.E. Halling s.n.; m, z. D.R. Sumstine s.n., 11 July 19??. All scale bars = 10 µm.
lanceolate, thin-walled, hyaline and lacking apparent contents, crowded and numerous. Pileipellis a hymeniderm
composed of one layer of versiform elements, terminal cells 37–144 × 13.0–32 µm, vesiculose, clavate,
cylindrical-clavate to fusoid cells (approx. 37–77 × 13.0–32 µm) intermixed by elongate-clavate to filiform
cystidioid elements (approx. 67–144 × 13.0–21 µm), apices typically rounded or rarely with a short papilla, thin-
Capítulo II - 69
walled and usually with brown intracellular contents. Stipitipellis a cutis of cylindrical hyphae, 3.0–21 µm wide,
thin-walled, hyaline, lacking apparent contents. Caulocystidia 32–64 × 11.0–27 µm, pyriform, clavate to
elongate-clavate or fusoid, apices typically rounded or rarely subacute, thin-walled, hyaline, often with brown
intracellular contents, scattered in fasciculate bundles, numerous. Clamp connections absent in all parts
examined.
Material examined BRAZIL: Paraná State, Foz do Iguaçu city, Parque Nacional do Iguaçu, 7 November 2010,
F. Karstedt FK1660 (SP); Rio de Janeiro State, Rio de Janeiro city, Jardim Botânico do Rio de Janeiro, Setor 36
C, 8 November 2002, A. A. Carvalho Jr. 20/02 (RB); São Paulo State, Palestina city, Fazenda Boa Vista, 10
December 2008, M. Capelari et al. MC4437 (SP); São Paulo city, Parque Estadual das Fontes do Ipiranga, 16
January 1987, D.N. Pegler 3714 [K(M) as P. pluvialis Singer]; 24 January 1987, D.N. Pegler et al. 3825 (SP as
P. pluvialis); 20 June 2006, F. Karstedt et al. FK679 (SP), FK680 (SP), FK681 (SP); 31 August 2008, L.A.S.
Ramos & F. Karstedt LASR25 (SP); Parque Estadual da Cantareira, Núcleo Engordador, 4 October 2008, F.
Karstedt & M. Capelari FK718 (SP); 4 December 2008, M. Capelari et al. MC4413 (SP); 16 February 2012, M.
Capelari & J.J.S. Oliveira MC4676 (SP). USA: Michigan, Montmorency city, NW Corner, 8 September 1969,
H.E.B. 15915 (NY); Missouri, Wayne city, Mingo National Wildlife Refuge, 20 September 2003, A.S. Methven,
Minnis 03-09-20-3 (SIU); New York, Albany city, July, leg, C.H. Peck (NYS #1760 – holotype); July 1921,
D.R. Sumstine s.n. (NY); Chautauqua city, 07 July 1942, D.R. Sumstine s.n. (NY); Nassau city, near Lake
Success, 5 June 1980, M. Morris s.n. (NY); Hamilton city, Blue Mountain Trail, Long Point, Raquette Lake, 27
July 1983, T.J. Baroni & R.E. Halling s.n. (NY); Pennsylvania, Bedford city, 11 July 19??, D.R. Sumstine s.n.
(NY); Centre city, State College Campus, 15 July 1936, M.B. Knauz s.n. (NY).
Fig. 10 Pileipellis elements of P. longistriatus. a. FK680; b. MC4431; c. FK681; d. Minnis 03-09-20-3; e. D.R. Sumstine s.n., July 1921;
f. M. Morris s.n.; g. T.J. Baroni & R.E. Halling s.n.; h. D.R. Sumstine s.n., 11 July 19??; i. H.E.B. 15915. All scale bars = 10 µm.
Habit and habitat Solitary, in small groups or subgregarious, on unidentified decaying wood or on rotting
wood of Acer, Pyrus and Quercus.
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Geographical distribution Pluteus longistriatus is a species apparently common in subtropical and temperate
areas of North and South America (Fig 1), and hitherto now, it has been recorded from Argentina (Singer 1959),
Brazil (Menolli and Capelari 2010; Gugliotta et al. 2012), Canada (Groves and Macrae 1963) and the USA (Peck
1878; 1885; Longyear 1904; White 1910; Kauffman 1915; 1918; Murrill 1917; Morse 1918; Sumstine 1918;
Singer 1959; Gilbertson et al. 1974; Branson 1989; Bates 2006; Menolli and Capelari 2010; Justo et al. 2011a;
b). Although the distribution of P. longistriatus in South America seems to be restricted (from Brazil it was
found only in Rio de Janeiro, São Paulo and Paraná States, and from Argentina in Prov. Salta), in the USA it is
widespread, especially in eastern region. Based on the materials examined herein and records from the
aforementioned references, P. longistriatus is certainly known in the USA from the states of Arizona,
Connecticut, Florida, Illinois, Kentucky, Massachusetts, Michigan, Missouri, New York and Pennsylvania.
There are probably also several unpublished herbarium records and other more obscure references that mention
P. longistriatus. The record of P. longistriatus from Canada is based on the collecting notes of Henry A. C.
Jackson from L'Islet, Prov. Quebec (Groves and Macrae 1963) and although there are no specimens preserved,
its occurrence seems to follow the distribution pattern along the northeastern USA.
Fig. 11 Holotype of P. longistriatus (C.H. Peck – NYS #1760). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia. d. Pileipellis elements.
e. Caulocystidia. All scale bars = 10 µm.
Discussion The occurrence of P. longistriatus is apparently correlated to summer in the northern hemisphere
(Peck 1878; Kauffman 1918) and also to autumn according to Longyear (1904), and it seems to be most
common during June to September. In the southern hemisphere, it can be found during the spring and summer,
commonly from October to February, but with some fructifications also occurring in winter during June/August.
Based on our samples, P. longistriatus can be considered as a well-delimited species regarding both
morphological and molecular characters. It is easily recognized usually by a pileus with a deeply sulcate-striate
margin and a hymenidermal pileipellis with heteromorphic elements. The only apparent difference between the
USA and Brazilian collections is the presence of a pale straw dissolved content in the pleurocystidia of the latter
(Fig. 9d– g). However, this could be considered an usual variation without geographical correlation since Singer
(1959) characterised all pleurocystidia (including from South America – Argentina) as without any intracellular
pigment.
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Acknowledgments
The authors thank the curators of ARAN, BR, F, ICN, INPA, K, MBM, MICH, NY, NYS, PACA, RSPF, SIU
and SP for managing the loans of their respective collections. We also thank Anderson Luiz dos Santos for
assistance with formatting the plates; Klei R. Sousa for assistance with the illustrations; Fernanda Karstedt and
Luiz Antonio S. Ramos as well as Walter J. Sundberg and Andrew S. Methven for variously collecting some
specimens and providing photographs used in this study; C.K. Pradeep for the Indian specimen; Flávia Ribeiro
Santos for helping with DNA isolation and PCR of some collections; Tatiane Asai for her helpful discussions.
N. Menolli Jr. and M. Capelari thank “Fundação de Amparo à Pesquisa do Estado de São Paulo” (FAPESP
grants 04/04319-2, 09/53272-2), “Coordenação de Aperfeiçoamento de Pessoal de Nível Superior” (CAPES PNADB grant 23038.000043/2010-43) and the “Conselho Nacional de Desenvolvimento Científico e
Tecnológico” (CNPq fellowships) for financial support. A. Justo acknowledges support from the National
Science Foundation grant DEB 0933081.
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CAPÍTULO III
Contributions to Pluteus (Pluteaceae, Agaricales) in Brazil: 21 taxa from the state of
Paraná including three new species
Nelson Menolli Jr., André A. R. de Meijer & Marina Capelari
Artigo a ser submetido para publicação na revista Nova Hedwigia.
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Contributions to Pluteus (Pluteaceae, Agaricales) in Brazil: 21 taxa from the state of
Paraná including three new species
Nelson Menolli Jr.*1,2, André A. R. de Meijer3 and Marina Capelari2
1
Instituto Federal de Educação, Ciência e Tecnologia de São Paulo, Campus São Paulo, CCT
/ Biologia, Rua Pedro Vicente 625, 01109-010 São Paulo, SP, Brazil
2
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902
3
PR-405 road, km 36, Guaraqueçaba, Paraná, PR, Brazil
*corresponding author: [email protected]
Abstract: To complement the knowledge of Pluteus in Brazil, we present a morphological
study of specimens collected by one of us in the state of Paraná between 1979 and 2011.
Amongst the total of 79 collections from Paraná, 17 collections were in too bad conditions for
identification and 18 were not encountered. All other were located at MBM and they
correspond to 21 species of Pluteus: 11 of sect. Celluloderma, seven of sect. Pluteus and three
of sect. Hispidoderma. Pluteus chusqueae, P. eludens, P. fusconigricans, P. homolae,
P. sapiicola and P. striatocystis are reported for the first time for Brazil. Pluteus
anomocystidiatus, P. crassus and P. neochrysaegis are newly proposed.
Keywords: Agaricales, Basidiomycota, Diversity, Taxonomy
Capítulo III - 79
Introduction
Pluteus Fr. (Pluteaceae, Agaricales) is a genus with worldwide distribution and includes ca.
300 species (Kirk et al. 2008). In the Neotropical region Pluteus species have been quite well
studied in many countries, including Argentina (Singer & Digilio 1952, Singer 1952, 1954a,
1956, 1959, 1962, 1969, Horak 1964, Raithelhuber 1991, Lechner et al. 2006), Belize (Pegler
1983), Bolivia (Singer 1959), Chile (Singer 1959, 1969, Horak 1964), Colombia (Singer
1973), Costa Rica (Ovrebo 1996), Cuba (Murrill 1911, Pegler 1983), Dominican Republic
(Justo et al. 2012), Ecuador (Singer 1978, Reid et al. 1981), French Guyana (Courtecuisse
1991), Grenada (Murrill 1917, Pegler 1983), Guadeloupe (Pegler 1983), Jamaica (Murrill
1911, Pegler 1983), Martinique (Pegler 1983) Mexico (Singer 1973, Vargas et al. 1993,
Candusso et al. 1994, Rodríguez & Guzmán-Dávalos 2000, 2001, Ramírez-Guillén &
Guzmán 2003, Pérez-Silva et al. 2006, Rodríguez et al. 2008, 2009, 2010), Panama (Singer
1989), Trinidad and Tobago (Baker & Dale 1951, Dennis 1953, Pegler 1983, Singer 1989),
and Venezuela (Dennis 1953, 1970, Singer 1962, Pegler 1983).
In Brazil, based on records in literature, about 70 taxa of Pluteus have been reported from ten
States: Amazonas (Hennings 1904a, Singer 1984, 1989, Singer & Aguiar 1986), Bahia
(Singer 1989), Minas Gerais (Rosa et al. 2003, Rosa & Capelari 2009), Mato Grosso
(Hennings 1900), Pará (Singer 1973, 1989), Paraná (Stijve & de Meijer 1993, Stijve 1995,
Meijer 2001, 2006, 2009, 2010), Rio de Janeiro (Singer 1956, 1959), Rio Grande do Sul (Rick
1907, 1919, 1930, 1938, 1961, Bresadola 1920, Singer & Digilio 1952, Singer 1954b, 1956,
1959, Pereira 1984, Raithelhuber 1991, Guerrero & Homrich 1999, Wartchow et al. 2004,
2006, Sobestiansky 2005, Drechsler-Santos et al. 2007), Rondônia (Singer 1959, Capelari &
Maziero 1988) and São Paulo (Hennings 1904b, Grandi et al. 1984, Pegler 1997, XavierSantos et al. 2004, Menolli & Capelari 2010, Menolli et al. 2010). However, many of these
records concern doubtful identifications or do not present descriptions for all species
mentioned, except for the works of Singer (1954b, 1956, 1959, 1973, 1989), Singer & Digilio
(1952), Wartchow et al. (2004, 2006), Menolli & Capelari (2010) and Menolli et al. (2010).
Revising the Pluteus collections from Brazil, we examined almost all specimens collected in
the state of Paraná by the second author between 1979 and 2011. Some of these materials
represent the 24 Pluteus taxa previously published from Paraná (Stijve & de Meijer 1993,
Meijer 2001, 2006, 2009, 2010). In these 32 years of study and observations, the collector
found about 34 species of Pluteus in Paraná (Meijer 2009), but many of the species
represented by a single basidiome only were not preserved, though annoted. Thus, with the
aim of complementing the study of Pluteus in Brazil, we present a re-examination of the
Capítulo III - 80
specimens collected by A.A.R. de Meijer to confirm the identifications previously published
and to present complete descriptions for the species recorded.
Material and methods
SAMPLING: The studied materials include specimens deposited at Herbarium MBM and acquired on numerous
random field trips, between 1979 and 2011, from different regions of Paraná. The herbarium acronyms follow
Thiers (2013). When necessary, type collections or additional materials from other Brazilian states were also
studied. The terminology of vegetation types is according to Veloso et al. (1991).
MORPHOLOGICAL STUDY: The macroscopic descriptions and accounts of habitat are exclusively based upon the
second author’s observations on the collections from Paraná, except for collections followed by an asterisk ‘*’
listed in the section ‘Additional collections examined’. In the latter case, the annotations made by the collector of
these specimens were also used to complement the macroscopic description. R means the radius of the pileus.
Colour codes employing the letters YR, A-F and NA(Y)M refer to Munsell (1975), Kornerup & Wanscher
(1978) and Küppers (1979), respectively.
For microscopic analyses, the dried material was wetted with 70% ethanol and then rehydrated in 5% KOH or
stained with Melzer’s reagent. All microscopic illustrations were made with the aid of a drawing tube. In a few
cases (e.g. P. cubensis and P. sapiicola) the better structures chosen for illustration are from Brazilian collections
outside Paraná. The notation [a/b/c] at the beginning of a set of basidiospores data is to be read as “(a)
basidiospores were measured from (b) basidiomata taken from (c) collections”. The basidiospores were
measured in lateral view, at least 20 basidiospores from each basidioma, and the terms denoting their shapes
follow Bas (1969). Q represents the range of the length/width or the quotient for all measured basidiospores, Qm
represents the average of all calculated Q values for all basidiospores measured and Lm (Wm) represents the
average of all measured basidiospores lengths (widths). For the pileipellis elements, the length of the pedicel was
included in the measurement of the spheropedunculate cells, and its length is also reported separately; the same
measurement criterion was applied when a long apical projection in some hymenial cystidium was observed.
Generic and infrageneric concepts follow Singer (1959, 1986) as complemented by Justo et al. (2011a, b).
Results
Among de Meijer’s Pluteus collections at MBM or those previously reported (Stijve & de
Meijer 1993, Meijer 2006, 2009, 2010), a total of 79 collections are listed in Table 1; 57 of
them being associated to the 24 taxa previously reported from Paraná [excluding
P. xylophillus var. tucumanensis that was reported by Stijve & de Meijer (1993) without
indication of material examined] and the other 22 represented by unreported species or
undetermined materials.
The confirmation of occurrence in the state of Paraná for some species was not possible
because some specimens were not preserved or because 35 collections were not found or are
in bad conditions for identification (Table 1). Thus, from this revision, we can not certify
Capítulo III - 81
from Paraná the records of P. cf. haywardii, P. laetifrons var. floridae Singer, P. cf.
polycystis, P. salicinus (Pers.) P. Kumm and P. cf. umbrinoalbidus. Moreover, we exclude the
records of P. chrysophlebius subsp. bruchii (Speg.) Singer, P. eucryphiae Singer,
P. eugraptus (Berk. & Broome) Sacc., P. cf. fluminensis, P. glaucus Singer, P. cf.
iguazuensis, P. laetifrons var. laetifrons (Berk. & M.A. Curtis) Sacc., P. aff. neophlebophorus
and P. cf. rimosoaffinis, because in this paper we redeterminate them (Table 1).
Among the remaining 43 collections of de Meijer, 21 species from Paraná were confirmed,
including six new records for Brazil and three species new to science (Table 1). Following the
infrageneric concept of Singer (1959, 1986) complemented by Justo et al. (2011a, b), sect.
Celluloderma is best represented in Paraná with 11 species, followed by sect. Pluteus and
sect. Hispidoderma, with seven and three species, respectively. The occurrence of more than
30 species from Paraná as previously reported by de Meijer (2009) seems plausible, as it was
not possible to re-examine many of his collections for this study. The importance of a broad
collecting effort allied to good conservation of the specimens must be emphasized. It is likely
that through continued collecting efforts in Paraná this number will be confirmed and even
exceeded.
Table 1: Pluteus collections from Paraná and notes about their identifications after re-examination.
Species name as reported by de
Meijer or as deposited at MBM
P. albostipitatus
P. cf. beniensis
P. chrysophlebius (Berk. & M.A.
Curtis) Sacc.
P. chrysophlebius subsp. bruchii
P. cubensis
P. cf. eucryphiae
P. eugraptus
P. fibulatus
P. cf. fluminensis
P. glaucus
Table 1, concluded.
de Meijer’s collector number# /
references
CUc-1557 (Meijer 2006)
MANa-2629 (Meijer 2006)
PAc-2744 (Meijer 2006)
REb-14.v.1992 (Meijer 2006)
CUb-31.xii.1991 (Meijer 2006)
MAe-2611 (unpublished data)
Identification after reexamination
Not found
P. albostipitatus
P. albostipitatus
Not found
Not found
P. sublaevigatus
PAa-1785 (Meijer 2006)
COa-4057 (Meijer 2006)
MAg-2.vi.1991 (Meijer 2006)
CUa-417 (Meijer 2006)
CUb-2085 (unpublished data)
CUh-2479 (Meijer 2006)
MANa-2631 (Meijer 2006)
FE-3300 (Meijer 2006)
SJf-3806 (Meijer 2006)
ANc-4215 (Meijer 2006)
CUa-284b (Meijer 2006)
CUa-284c (Meijer 2006)
CUa-208c (Meijer 2006)
SJa-9.ii.1992 (Stijve & de Meijer 1993,
Meijer 2006)
Not found
P. crassus
Not found
P. rimosellus
P. glaucotinctus
Insufficient for identification
P. cubensis
P. striatocystis
P. rimosellus
P. hylaeicola
P. chusqueae
P. fibulatus
Not found
P. fibulatus
P. riograndensis
P. riograndensis
P. riograndensis
Not found
Species name as reported by de
Meijer or as deposited at MBM
P. glaucus
P. aff. harrisii
P. cf. haywardii
P. cf. iguazuensis
P. laetifrons var. laetifrons
P. laetifrons var. floridae
P. aff. neophlebophorus
P. cf. polycystis
P. pulverulentus var. pseudonanus
Singer
de Meijer’s collector number# /
references
MAh-1670 (Meijer 2006)
CUb-2562 (Meijer 2006)
CUb-2596 (Meijer 2006)
SJa-9.ii.1992 (Meijer 2006)
Capítulo III - 82
Identification after reexamination
Not found
P. glaucotinctus
P. glaucotinctus
Not found
P. glaucotinctus
P. albostipitatus
Not found
P. crassus
Not found
P. eludens
P. eludens
Not found
Not found
Not found
Not found
P. sapiicola
P. cf. puttemansii
MAg-3717 (Meijer 2010)
P. harrisii
P. cf. rimosoaffinis
MAg-11.iv.1992 (Meijer 2006)
Not found
CUi-2196 (Meijer 2006)
P. riograndensis
CUb-2552a (Meijer 2006)
P. sapiicola
CUb-2552b (Meijer 2006)
Not found
P. salicinus (Pers.) P. Kumm.
MAc-5.xii.1992 (Stijve & de Meijer 1993 Not found
as P. glaucus, Meijer 2006)
P. aff. thomsonii
P. thomsonii
P. cf. umbrinoalbidus
not preserved (Meijer 2006)
Not preserved
P. xylophillus
CUa-160 (unpublished data)
P. xylophillus
CUa-160b (unpublished data)
P. xylophillus
SJf-4023 (Meijer 2006, 2009)
P. xylophillus
SJf-4086 (Meijer 2006, 2009)
P. xylophillus
P. cf. angustisporus
ANc-4165 (Meijer 2006, 2009)
P. xylophillus
Pluteus spp.
P. riograndensis
P. fusconigricans
P. homolae
P. neochrysaegis
P. neochrysaegis
CUb-2549 (umpublished data)
ALAa-3210 (unpublished data)
JSa-3252 (unpublished data)
JSa-3253 (unpublished data)
JSb-3258 (unpublished data)
JSb-3259 (unpublished data)
P. homolae
DN-3378 (unpublished data)
P. anomocystidiatus
COa-3628 (unpublished data)
COa-3930 (unpublished data)
ANc-4435 (unpublished data)
P. anomocystidiatus
GUf-4480 (unpublished data)
P. riograndensis
# = codes that precede the collector numbers represent the collecting area as previously used in Meijer (2006,
2009, 2010).
Taxonomy
Pluteus albostipitatus (Dennis) Singer, Lloydia 21: 240. 1959.
Capítulo III - 83
Fig. 1
≡ P. spilopus var. albostipitatus Dennis, Bull. Soc. Mycol. Fr. 69: 195. 1953.
= P. albostipitatus var. poliobasis Singer, Beih. Sydowia 7: 64. 1973.
= P. phaeoleucus E. Horak, Bull. Jard. Bot. Belg. 47: 89. 1977.
= P. melanopotamicus Singer, Fieldiana, Bot. 21: 96. 1989.
= P. densifibrillosus Menolli & Capelari, Mycologia 102: 698. 2010.
PILEUS 23–45 mm diam, at first strongly convex, then plano-convex, finally applanate,
sometimes subumbonate, hygrophanous or not, translucently striate < 2/3 R, dark brown
(6-7[F]4-6) to yellowish brown at centre, greyish brown (5D3) to brown (6-7[E]3-4) between
fibrils, surface smooth and not fibrillose (most collections), or entirely radially innate
fibrillose on paler background. LAMELLAE close to moderately close, free and up to 1.5 mm
from stipe, subventricose, < 5 mm broad, at first white then becoming pink. STIPE 22–60 ×
2.5–5(–7 at base) mm, central, cylindrical or slightly tapering upwards, base clavate or equal,
solid, at first (young stipe) entirely pale brownish grey (5C2), finally white but the lower part
often very pale greyish brown striate, basal mycelium pure white. FLESH pure white, < 2 mm
thick in centre and < 0.5 mm thick above midpoint of lamellae. ODOUR indistict. TASTE not
recorded. SPORE PRINT pink. BASIDIOSPORES [120/4/4] 6.2–8.7 × 5.0–7.5 µm [Q = 1.00–1.24(–
1.36); Qm = 1.14; Lm = 7.1 µm; Wm = 6.2 µm], globose to broadly ellipsoid, rarely ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 35 × 9.0 µm, urniform, thinwalled, four-spored. PLEUROCYSTIDIA 44–81 × 12.5–32 µm, slightly lageniform to slightly
utriform, sometimes (slightly) clavate, with rounded, subcapitate or truncate apex, some with
an apical constriction like a neck and forming two rounded or (only in FK1664) two to four
well-developed prongs at apex or with several small finger-like protuberances, colourless,
hyaline, thin- to slightly thick-walled, sometimes wall thickened up to 1.0 μm wide,
moderately numerous. CHEILOCYSTIDIA 33–72(–102) × (8.7–)10.0–17.5 µm, mostly clavate
with a well-developed pedicel at the base or (only in MANa-2629) slightly ventricose to
lageniform, colourless, hyaline, thin-walled, abundant and mostly in fascicules. PILEIPELLIS a
cutis composed of cylindrical, thin-walled hyphae, 10.0–13.0 µm diam, with rounded apex or
gradually attenuated to obtuse apex (3.0–4.0 µm diam), mostly with evenly dissolved
brownish intracellular pigment. CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: Solitary, in groups of two or subgregarious, on decayed
branches and trunks of dicotyledonous wood. In Paraná it was found in dense and mixed
ombrophilous lowland to montane and alluvial forests, from sea level to 850 m, fruiting in
November, April and May. Considering the records from other countries it seems to occur
Capítulo III - 84
year round (Menolli et al. 2013). Tropical to subtropical occurrence is known from several
countries in the Americas, Africa and Asia (Dennis 1953, Singer 1956, 1959, 1973, 1989,
Horak 1964, 1977, Horak & Heinemann 1978, Reid et al. 1981, Pegler 1983, Courtecuisse
1991, Raithelhuber 1991, Stijve & de Meijer 1993, Pegler 1997, Rodríguez & GuzmánDávalos 2000, 2001, Meijer 2001, 2006, Wartchow et al. 2006, Menolli & Capelari 2010,
Menolli et al. 2010, 2013, Yang & Bau 2010, Justo et al. 2011a, b). Type locality: Trinidad,
Diego Martin.
Fig. 1. Pluteus albostipitatus. a–c. basidiospores. d–f. pleurocystidia. g–h. cheilocystidia. a, d, g. FK1664.
b, e, h. MANa-2629. c, f, i. PAc-2744. Bars = 10 µm.
Capítulo III - 85
MATERIAL EXAMINED: BRAZIL, Paraná: Antonina, Reserva Natural do Rio Cachoeira, 24 May 2003, A.A.R. de
Meijer ANc-4219 (MBM); Foz do Iguaçu, Parque Nacional do Iguaçu, 7 Nov 2010, F. Karstedt FK1664 (SP);
Mandirituba, 20 April 1993, A.A.R. de Meijer MANa-2629 (MBM); Paranaguá, Mundo Novo, 27 May 1993,
A.A.R. de Meijer PAc-2744 (MBM).
COMMENTS: The wide morphological variation observed in P. albostipitatus was recently
discussed by Menolli et al. (2013) who included many synonyms under this name. However,
the authors concluded that the same morphological species concept probably includes more
than a single phylogenetic species. The morphological variation of pleurocystidium shape can
be verified also in the collections from Paraná, like in FK1664 (Fig. 1d) that has
pleurocystidia with slightly thickened to thickened walls and well-developed prongs (like
those of P. melanopotamicus, see Menolli et al. 2013) or like MANa-2629 (Fig. 1e) and PAc2744 (Fig. 1f) that have thin-walled pleurocystidia with rounded or truncate apex,
respectively. Regarding the shape of cheilocystidia it was also verified a variation in the
collections from Paraná, e.g. in MANa-2629 (Fig. 1h) that has slightly ventricose to
lageniform cheilocystidia (like those of P. densifibrillosus, see Menolli & Capelari 2010 and
Menolli et al. 2013) or in PAc-2744 (Fig. 1i) and FK1664 (Fig. 1g) that have mostly clavate
cheilocystidia with a well-developed pedicel.
Although P. albostipitatus was classified originally in sect. Hispidoderma (Singer 1959),
recent molecular studies of Pluteus phylogeny confirmed its position in sect. Pluteus (Menolli
et al. 2010, 2013, Justo et al. 2011a, b).
Pluteus cf. angustisporus Singer, Lloydia 21: 206. 1959.
Fig. 2
PILEUS 32 mm diam, convex, subumbonate, brown, surface smooth. LAMELLAE very close,
free and 1.0 mm remote from stipe, ventricose, 5 mm broad, pure white (probably an imature
specimen). STIPE 35 × 4 mm, central, cylindrical, solid, whitish. FLESH pure white, < 4 mm
thick near stipe. ODOUR slightly fungoid. TASTE fully mild. SPORE
PRINT
not obtained.
BASIDIOSPORES [20/1/1] 5.0–6.2 × 3.7–4.3 µm (Q = 1.35–1.68; Qm = 1.51; Lm = 5.6 µm; Wm
= 3.8 µm), ellipsoid to elongate, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA
26 × 7 µm, clavate, thin-walled, four spored. PLEUROCYSTIDIA 50–77 × 16.2–24 µm, fusoidventricose, colourless, hyaline, metuloids of the Cervinus-type with four to six apical or
lateral prongs, thick-walled (< 1.5 µm wide), abundant to very abundant. CHEILOCYSTIDIA
thin-walled, colourless, hyaline, apparently dimorphic: I) 52–111 × 11.0–15.0 µm, clavate
with slender pedicel, crowded; II) 32–55 × 13.7–45 µm, short-clavate or pyriform to
Capítulo III - 86
spheropedunculate, scarce. PILEIPELLIS a cutis composed of parallel chains of cylindrical,
thin-walled hyphae, individual terminal elements up to 125 × 8.0 µm, mostly with rounded
apex and brown intracellular pigment, sometimes almost colourless or with condensed brown
pigment. CLAMP CONNECTIONS absent.
HABIT, HABITAT, AND DISTRIBUTION: Solitary. In Paraná it was found in mixed ombrophilous
forest, at the base of a dicotyledonous stub at 900 m above sea level. Fruiting in October (this
record) and in April (type collection, Singer 1959). Menolli & Capelari (2013) listed some
additional old Brazilian collections most likely represent P. angustisporus. Type locality:
Bolivia, Prov. Madre de Dios.
MATERIAL
EXAMINED:
BRAZIL,
Paraná:
São
José
dos
Pinhais,
Roça
Velha,
31
Oct
2002,
A.A.R. de Meijer SJf-4113 (MBM).
Fig. 2. Pluteus cf. angustisporus (SJf-4113). a. basidiospores. b. pleurocystidia. c. cheilocystidia. Bar = 10 µm.
COMMENTS: Pluteus angustisporus is morphologically close to P. xylophilus and P. harrisii.
However, P. angustisporus has remarkably narrower basidiospores than P. harrisii, and when
compared to P. xylophilus the absence of pleurocystidia of the Magnus-type in
P. angustisporus is notable. In addition, Singer (1959) also reported for P. angustiporus the
occurrence of polymorphic pileipellis elements and cheilocystidia; in Paraná collection the
former could not be recovered, but the latter were indeed polymorphic by the second author
when he examined the fresh specimen, although much longer (50–115 µm long) then found
by Singer (1959) in the type (16.5–44 µm long). During the re-examination of SJf-4113 the
first author only found a single short cheilocystidia (Fig. 2c). The presence of narrow
basidiospores and the pleurocystidium features are characters that fit within the concept of
Capítulo III - 87
P. angustiporus. However, we prefer to maintain the collection SJf-4113 as uncertain
determination due to the failure to recover some features during the re-examination and also
due to the occurrence of very long cheilocystidia observed by the second author. Recently,
Menolli & Capelari (2013) tentatively attributed this name to some old Brazilian collections
studied by J. Rick and P. Hennings. Pluteus angustisporus belongs to sect. Pluteus.
Pluteus anomocystidiatus Menolli & de Meijer sp. nov.
Figs. 3–4
HOLOTYPUS: BRAZIL, Paraná: Diamante do Norte, Estação Ecológica do Caiuá, 18 Mar 1996, A.A.R. de Meijer
DN-3378 (MBM).
ETYMOLOGY: The name refers to the shape of the pleuro- and cheilocystidia that is unusual for Pluteus.
PILEUS 13–25 mm diam, strongly convex to plano-convex, subumbonate, not translucently
striate or only so at extreme margin, evenly dark brown (6F5, 8F4), surface punctate, smooth.
LAMELLAE close, free, ventricose, 3–4 mm broad, at first white then becoming orange white
(5A2) to greyish orange (5B3), entire edge strongly contrasting dark brown (6F4) or greyish
brown (6F3). STIPE 22–30 × 1.7(middle)–3(base) mm, central, slightly attenuated from base
towards the apex or evenly cylindrical, base clavate, solid becoming narrowly fistulose,
entirely dense dark brown (6F4) punctate-flocculose on pale greyish white ground colours.
FLESH greyish yellow (4B3), < 1.3 mm thick in centre and < 0.5 mm thick above midpoint of
lamellae. ODOUR indistinct. TASTE and
SPORE PRINT COLOUR
not recorded. BASIDIOSPORES
[40/2/2] (3.7–)4.3–5.0(–5.6) × (3.1–)3.7–4.3(–5.0) µm [Q = 1.00–1.16(–1.39); Qm = 1.15; Lm
= 4.7 µm; Wm = 4.1 µm], globose to subglobose, rarely broadly ellipsoid or ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 26–31 × 7.5–10.0 µm, clavate,
thin-walled, four-spored. PLEUROCYSTIDIA 29–33 × 16.2–20 µm, clavate, vesiculose or
spheropedunculate, thin-walled, filled with condensed or sometimes evenly dissolved
brownish intracellular pigment, scarce. CHEILOCYSTIDIA 50–116 × 12.5–25 µm, elongatefusiform or filiform, sometimes clavate, apices typically rounded or slightly obtuse and
gradually attenuated to apex 5.0 µm diam, filled with condensed or sometimes evenly
dissolved brownish intracellular pigment but some also dissolved, crowded. LAMELLAR EDGE
sterile. LAMELLAR TRAMA inverse, approx. 31 µm wide, composed of thin-walled hyphae up
to 7.5 µm diam, hyaline. PILEUS CONTEXT undifferentiated, composed of thin-walled hyphae
up to 7.5 µm diam, hyaline. PILEIPELLIS a hymeniderm composed of a single layer of clavate,
subglobose or spheropedunculate cells, 32–55 × 18.7–32 µm, with a short to medium pedicel
(2.5–11.2 µm long), intermixed with elongate-fusiform or elongate-clavate cystidioid
Capítulo III - 88
elements, 75–112 × 15–25 µm, both types thin-walled and filled with condensed or sometimes
evenly dissolved brownish intracellular pigment. CAULOCYSTIDIA 47–140 × 12.5–17.5 µm,
elongate-fusiform or elongate-clavate, filled with condensed or sometimes evenly dissolved
brownish intracellular pigment, very numerous. CLAMP CONNECTIONS absent.
HABITAT,
HABIT AND DISTRIBUTION:
Solitary or subgregarious. Only known from Paraná
where it was found in dense ombrophilous lowland and seasonal semi-deciduous submontane
forests, on decayed wood of dicotyledonous, from sea level to 300 m, fruiting in March.
MATERIAL EXAMINED: BRAZIL, Paraná: Diamante do Norte, Estação Ecológica do Caiuá, 18 Mar 1996, A.A.R.
de Meijer DN-3378 (MBM – Holotype); Antonina, Reserva Natural do Rio Cachoeira, 7 Mar 2009, A.A.R. de
Meijer ANc-4435 (MBM).
Fig. 3. Pluteus anomocystidiatus (DN-3378 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia.
Bars = 10 µm.
Capítulo III - 89
Fig. 4. Pluteus anomocystidiatus (DN-3378 – holotype). a. pileipellis elements. b. caulocystidia. Bars = 10 µm.
COMMENTS: Considering the structure of the pileipellis, P. anomocystidiatus is classified in
sect. Celluloderma (subsect. Mixtini according to Singer 1959, 1986). Among the species
known in that group, the combination of a pileipellis containing both elongated and short
clavate-spheropedunculate elements with the uniformly dark lamellar edges is observed in
very few species, viz. P. eludens, P. eugraptus and P. multiformis Justo, A. Caball. &
G. Muñoz (Singer 1956, 1959, Pegler 1977, 1986, Justo et al. 2011c). Pluteus
anomocystidiatus clearly differs from these species in the shape and size of pleuro- and
Capítulo III - 90
cheilocystidia. The presence of short spheropedunculate pleurocystidia combined with very
long elongate-fusiform cheilocystidia and the other characters mentioned above are our reason
to propose this as a new species.
Pluteus chusqueae (E. Horak) Menolli, stat. nov.
Fig. 5
≡ P. psychriophorus var. chusqueae E. Horak, Nova Hedwigia 8: 189. 1964.
≡ P. eugraptus var. chusqueae (E. Horak) Singer, Beih. Nova Hedwigia 29: 157. 1969.
PILEUS 13 mm diam, convex to plano-convex, when fresh translucently striate < ½ R, (dark)
brown (6EF5, 6F5) to dark yellowish brown (5.5F6), surface smooth. LAMELLAE moderately
close, free, lanceolate, < 2.2 mm broad, at first white and then becoming orange grey (6C2),
edge pale brown (sometimes very faint). STIPE 15 × 1.2 mm, central, cylindrical, solid, white
to brownish grey (6C2) with darker base, surface smooth, slightly dark brown fibrillose on
pale background, more strongly fibrillose towards the base. ODOUR indistinct. FLESH COLOUR,
TASTE
and SPORE PRINT COLOUR not recorded. BASIDIOSPORES [20/1/1] 5.0–6.2(–6.8) × (4.3–)
5.0–6.2 µm [Q = 1.00–1.12(–1.24); Qm = 1.06; Lm = 5.7 µm; Wm = 5.3 µm], globose to
subglobose, rarely broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate.
BASIDIA 25 × 8.5 µm, ventricose, thin-walled, four-spored. PLEUROCYSTIDIA up to 87 × 30
µm, mostly lageniform with tapering neck, apex obtuse, sometimes subcapitate up to 10 µm
broad at the apex, thin-walled, filled with brownish intracellular pigment, scarce.
CHEILOCYSTIDIA 38–70 × 11.0–27 µm, broadly clavate to vesiculose or slightly utriform, thinwalled, filled with brownish intracellular pigment, crowded. PILEIPELLIS a hymeniderm
composed of a single layer of versiform elements, 30–180 × 13.7–33 µm, vesiculose, clavate,
slightly utriform or spheropedunculate (approx. 30–80 × 15–33 µm) intermixed by elongatefusiform or elongate-clavate to filiform cystidioid elements (80–180 × 13.7–26 µm), apices
typically rounded or attenuated to slightly obtuse (3 µm diam), usually with a short mucro,
with either condensed or evenly dissolved brownish intracellular pigment. CAULOCYSTIDIA
50–60 × 8.7–12.5 µm, versiform or elongate-fusiform to elongate-clavate, filled with evenly
dissolved brownish intracellular pigment, numerous. CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: Solitary. In Paraná it was found in mixed ombrophilous
montane forest, on decayed trunks of dicotyledonous at 900 m above sea level. Fruiting in
April (this record) and in March (type collection, Horak 1964). Type locality: Argentina,
Prov. Neuquen.
Capítulo III - 91
Fig. 5. Pluteus chusqueae (COa-3748). a. basidiospores. b. cheilocystidia. c. pileipellis elements.
d. caulocystidia. Bars = 10 µm.
MATERIAL EXAMINED: BRAZIL, Paraná: Colombo, Embrapa Florestas, 10 Apr 2000, A.A.R. de Meijer &
M.A.L. de A. Amazonas COa-3748 (MBM).
COMMENTS: Horak (1964) described P. psychriophorus var. chusqueae mainly based on its
growth on bamboo substrates (Chusquea culeou E. Desv.). The longer cystidioid elements in
the pileipellis as described for this variety [(45–)70–100 × (25–)27–34 µm] distinguish it
clearly from the type variety, which according to Singer (1956) has much smaller pileipellis
Capítulo III - 92
cells (39–48 × 13–16.3 µm). Singer (1969) proposed a new combination in P. eugraptus var.
chusqueae. But according to the redescription of the type made by Justo et al. (2011c),
P. eugraptus var. eugraptus has shorter pileipellis cells (20–70 × 10–25 µm). Singer (1956,
1959) reported pileipellis elements up to 140 µm long for collections of P. eugraptus var.
eugraptus from South America, which probably represent P. chusqueae.
The presence of a hymeniderm of short cells mixed with cystidioid elements clearly put
P. eugraptus var. eugraptus and P. psychriophorus var. psychriophorus in the sect.
Celluloderma (subsect. Mixtini according to Singer 1959, 1986), unlike P. chusqueae that is
better classified in sect. Hispidoderma and here considered different from P. eugraptus and
P. psychriophorus. This sole character justifies the proposition of the new status for
P. eugraptus var. chusqueae.
Regarding the Paraná collection there are minor differences from the description presented by
Horak (1964): our material has a pale brown lamellar edge and was collected on
dicotyledonous trunks. However, in Pluteus these characters can be considered of
intraspecific variations. For instance, in P. eludens the basidiomata may have either a darkbrown or a concolourous lamellar edge (Justo et al. 2011c), and P. harrisii (= P. cervinus var.
bambusinus Baker & Dale) has been reported both from a bamboo stool (Singer 1956, 1959,
Pegler 1983) as from dead and rotting dicotyledonous wood.
Pluteus crassus Menolli & de Meijer sp. nov.
Figs. 6–7
HOLOTYPUS: BRAZIL, Paraná: Colombo, Embrapa Florestas, 21 Mar 2001, A.A.R. de Meijer & M.A.L. de A.
Amazonas COa-4057 (MBM).
ETYMOLOGY: The name refers to the wall thickness of the pleuro- and cheilocystidia that is unusual for species
of sect. Celluloderma.
PILEUS 25–39 mm diam, plane with deflexed margin and centre usually depressed,
translucently striate < ½ R, dry and slightly hygrophanous, centre yellowish brown (5F5), and
elsewhere light brown (5D5) to olive brown/yellowish brown (4.5E8), minutely innatelyfibrillose at the centre and smooth elsewhere. LAMELLAE moderately close, free to narrowly
adnexed, subventricose, < 5.5 mm broad, white. STIPE 30–52 × 2–4(apex)–5(base) mm,
central, cylindrical, base narrowly clavate, yellowish white (3A2), base dull yellow (3B3) to
dark yellow (3B4), surface translucently striate, dry, basal third minutely light brown to olive
brown punctate, elsewhere glabrous, basal mycelium a dense pure white. FLESH in fresh
Capítulo III - 93
pileus olive (3D3) to olive brown brown (4F3), drying white, < 2 mm thick near stipe and
< 0.5 mm thick above midpoint of lamellae. ODOUR indistinct. TASTE and
SPORE PRINT
not
recorded. BASIDIOSPORES [40/2/2] 5.0–6.2 × (4.3–)5.0–5.6 µm [Q = 1.00–1.16(–1.30);
Qm = 1.09; Lm = 5.5 µm; Wm = 5.1 µm], globose to subglobose, rarely broadly ellipsoid,
inamyloid, hyaline, smooth, thick-walled. BASIDIA 20–26 × 7.5–8.7 µm, clavate, thin-walled,
four-spored. PLEUROCYSTIDIA 45–80 × 13.7–30 µm, mostly clavate or sometimes narrowly
Fig. 6. Pluteus crassus (COa-4057 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis
elements. Bars = 10 µm.
Capítulo III - 94
lageniform, usually with dissolved light brown intrecellular pigment and an apical mucilage,
thin-walled or frequently upper part up to 1.2 µm thick wall, abundant. CHEILOCYSTIDIA 44–
110 × 11.2–24 µm, clavate to vesiculose or slightly ventricose to lageniform or utriform, neck
either slightly attenuated to apex or subcapitate, at apex 6.0–11.0 µm broad, usually with
dissolved light brown intracellular pigment and an apical mucilage strongly adhered to the
wall, thin-walled or frequently wall up to 1.2 µm thick, thickened all over or only at upper
part, crowded. LAMELLAR
EDGE
sterile. LAMELLAR
TRAMA
inverse, up to 44 µm wide,
composed of thin-walled, hyaline hyphae, 3.7–10.0 µm diam, hyaline, frequently with
oleiferous hyphae up to 7.5 µm diam. PILEUS CONTEXT undifferentiated, approximately 81 µm
thick, composed of thin-walled, hyaline hyphae, 3.7–10.0 µm diam, frequently with oleiferous
hyphae up to 7.5 μm diam. PILEIPELLIS a hymeniderm up to 75 µm thick, composed of one
layer of thin-walled and clavate, vesiculose or subglobose cells, 23–41 × 17.5–24 µm, usually
non-pedicellate but sometimes with a short pedicel (2.5–8.7 µm long), with evenly dissolved
brown intracellular pigment. CLAMP CONNECTIONS absent.
HABIT,
HABITAT AND DISTRIBUTION:
Solitary or in groups of two. Only known from Paraná
where it was found in dense and mixed ombrophilous alluvial to montane forests, on decayed
branch of dicotyledonous, from sea level to 900 m. Fruiting only in March and May.
Meijer ANc-4203 (MBM). Colombo, Embrapa Florestas, 21 Mar 2001, A.A.R. de Meijer & M.A.L. de
A. Amazonas COa-4057 (MBM – Holotype).
Fig. 7. Pluteus crassus (COa-4057 – holotype). a–b. pleurocystidia. c–d. cheilocystidia. Bars = 10 µm.
Capítulo III - 95
COMMENTS: Considering the structure of the pileipellis, P. crassus is classified in sect.
Celluloderma. However, differently from any other species of this section, P. crassus has
thick-walled pleuro- and cheilocystidia.
The presence of thick-walled cystidia is commonly associated to species of sect. Pluteus
(Singer 1959, Justo et al. 2011a). However, recently Pradeep et al. (2012) confirmed the
position of P. chrysaegis (Berk. & Broome) Petch in sect. Hispidoderma and thus
demonstrated the occurrence of thick-walled cystidia in species of that section (see also
comments under P. neochrysaegis). The discovery of P. crassus in sect. Celluloderma shows
that the character of thick-walled cystidia has originated independently at least three times
during evolution in the genus.
Pluteus cubensis (Murrill) Dennis, Bull. Soc. Mycol. Fr. 69: 155. 1953.
Fig. 8
PILEUS 12–40 mm diam, broadly convex-umbonate, becoming applanate, strongly radially
silky-fibrillose, fibrils very dark brown (6F3) to nearly black on dark yellowish brown
(5F4-5) ground colour, centre finely granulose-scaly, fibrils appressed and dissociating in the
marginal zone where showing the white flesh, margin short-striate. LAMELLAE close, free and
remote from stipe, pinkish, with two to three series of lamellulae. STIPE 18–50 × 1–4 mm,
central, cylindrical, sometimes slightly broadened at base, solid, entirely pure white, surface
smooth, sometimes slightly pruinose at the base. ODOUR,
PRINT COLOUR
TASTE, FLESH COLOUR
and
SPORE
not recorded. BASIDIOSPORES [140/7/7] 5.0–7.5(–8.7) × 5.0–6.2(–8.7) µm
[Q = 1.00–1.24(–1.50); Qm = 1.14; Lm = 6.8 µm; Wm = 6.0 µm], globose to broadly
ellipsoid, rarely ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA
25–31(–37) × 7.5–10.0 µm, versiform to clavate, sometimes more slender, thin-walled, fourspored. PLEUROCYSTIDIA 43–55(–77) × (12.5–)13.7–20(–22) µm, subclavate to clavate,
sometimes elongate-ventricose, colourless and hyaline, thin-walled, not numerous, deeply
rooting and difficult to see. CHEILOCYSTIDIA (20–)31–47(–56) × (8.7–)10.0–21(–25) µm,
mainly clavate to clavate-pedicellate, colourless and hyaline, thin-walled, in fascicles and
abundant, in some collections scarce. LAMELLAR
EDGE
heteromorphous. LAMELLAR
TRAMA
inverse, up to 31 µm wide, composed of thin-walled, hyaline hyphae, 2.5–8.7 µm diam.
PILEUS
CONTEXT
undifferentiated, approximately 120 µm thick, composed of thin-walled,
hyaline hyphae, 2.5–8.7 µm diam. PILEIPELLIS a cutis, approximately 50 µm thick, composed
of thin-walled hyphae, 7.5–12.0 µm diam, with a few elongated, ascendant, conspicuous
Capítulo III - 96
subradial elements, with rounded to subacute apex (up to 5.0 µm diam) and evenly dissolved
brownish intracellular pigment. CLAMP CONNECTIONS absent.
HABIT, HABITAT, AND DISTRIBUTION: Solitary, in groups of two or subgregarious. In Paraná it
was found in mixed ombrophilous montane forest, on decayed branches of dicotyledonous
wood, from 800 to 900 m above sea level, fruiting in April. Considering the other materials
herein studied and previous records from other countries, it seems to fruit year round.
Neotropical distribution: Cuba (Murrill 1911), Trinidad (Dennis 1953), Argentina (Singer
1956, 1959), Martinique (Pegler 1983) and Brazil (Stijve and de Meijer 1993, Pegler 1997,
Meijer 2006). Type locality: Cuba, Santiago de las Vegas.
COLLECTION EXAMINED: BRAZIL, Paraná: Mandirituba, 20 Apr 1993, A.A.R. de Meijer MANa-2631 (MBM).
ADDITIONAL COLLECTIONS EXAMINED: BRAZIL, Rondônia: Ji-Paraná, Reserva Biológica do Jarú, Margem
direita do Rio Ji-Paraná, 11 May 1987, M. Capelari et al. MC1367 (SP); near Ilha 7 de setembro, 14 May 1987,
M. Capelari et al. MC1526 (SP); São Paulo: Iporanga, PETAR, Núcleo Ouro Grosso, 23 Mar 2007, C. Puccinelli
et al. CP233 (SP)*; Mogi Guaçu, Reserva Biológica de Mogi Guaçu, Fazenda Campininha, 30 Jan 1987, D.N.
Pegler et al. 3887 (K); CUBA, Santiago de las Vegas, 29 Aug 1904, F.S. Earle 155 (NY – Holotype);
TRINIDAD, Vicinity of Simla Research Station, 3 Jan 1984, T.J. Baroni 4461 (NY).
COMMENTS: Pluteus cubensis has a wide morphological variation especially regarding the
basidiospores shape and size. Also Singer (1956) considered P. cubensis a variable and
deceptive species of wide distribution specially in Central and South America. The present
work provides for the first time a complete description and revision of the Brazilian
collections of P. cubensis.
Murrill (1911) described P. cubensis with subglobose basidiospores, as also observed in the
materials studied by Dennis (1953), Singer (1959) and Pegler (1983). Pegler (1983)
characterized the basidiospores as almost globose like those observed in some Brazilian
collections herein studied, which vary from globose to broadly ellipsoid. The cheilo- and
pleurocystidia observed are similar to those reported by Dennis (1953), Singer (1956, 1959)
and Pegler (1983). The granular incrustation or the nodulose excrescenses in the
pleurocystidia mentioned by Singer (1956, 1959) were not observed by us.
The re-examination of the type material shows that basidiospores (Fig. 8a) are preponderantly
broadly ellipsoid and rarely subglobose or ellipsoid [20/1/1] 7.5–8.7(–10.0) × (5.0–)6.2–7.5
(–8.7) µm [Q = (1.15–)1.16–1.21(–1.50); Qm = 1.23; Lm = 8.0 µm; Wm = 6.6 µm]. The
pleurocystidia were not recovered and only two cheilocystidia (Fig. 8l) measuring 26–29 ×
6.2–10.0 µm were seen.
Capítulo III - 97
Dennis (1953) considered P. fibrillosus Murrill a synonym of P. cubensis. Smith & Stuntz
(1958) reported incrusted pileipellis elements in P. fibrillosus, and this character has never
been reported for P. cubensis. Banerjee & Sundberg (1993) confirmed the presence of
incrusted pileipellis, nevertheless they considered P. cubensis and P. fibrillosus as synonyms,
although mistakenly treating P. cubensis a synonym of P. fibrillosus and not the inverse.
Since then, the relationship between P. cubensis and P. fibrillosus has not been discussed.
Fig. 8. Pluteus cubensis. a–e. basidiospores. f. basidia. g–j. pleurocystidia. k–o. cheilocystidia. a, k. F.S. Earle
155 (holotype). b, g, l. MC1367. c, h, m. D.N. Pegler et al. 3887. d, i, n. T.J. Baroni 4461. e, f, j, o. CP233. Bars
= 10 µm.
Capítulo III - 98
Based on the re-examination of the P. fibrillosus holotype [U.S.A., Louisiana, New Orleans,
F.S. Earle 129, 8 Sept. 1908 (NY)] we consider them as distinct species due to the incrusted
pileipellis elements (Fig. 9a) and, as mentioned by Singer (1956), due to the presence of
cystidioid and versiform elements present in the pileipellis, whilst in P. cubensis the
pileipellis is a typical cutis. Furthermore, the pleuro- and cheilocystidia (Fig. 9b–c) are
slightly longer [(44–)50–95 × (12.5–)15.0–27 µm and 47–59 × (10.0–)12.5–21 µm,
respectively] than those of P. cubensis and the cheilocystidia are slightly ventricose like the
pleurocystidia and not clavate to clavate-pedicellate as in P. cubensis. Finally, the
basidiospores of the type material of P. fibrillosus are preponderantly broadly ellipsoid and
occasionally globose [20/1/1] 6.2–7.5 × 5.0–6.2 µm [Q = (1.00–)1.21–1.24; Qm = 1.19;
Lm = 6.5 µm; Wm = 5.5 µm].
Fig. 9. Pluteus fibrillosus (F.S. Earle 129 – holotype). a. pileipellis elements. b. pleurocystidia. c. cheilocystidia.
Bars = 10 µm.
After re-examination of collections identified as P. cubensis from Paraná and other Brazilian
States, a few corrections and considerations must be made. Some collections listed by Meijer
(2006) actually represent P. rimosellus (CUa-417 and ANc-4215), P. striatocystis (FE-3300)
or still another unidentified species (SJf-3806) characterized mainly by globose to subglobose
basidiospores, capitate pleurocystidia frequently with incrusted wall and a pileipellis like a
cutis. Other collections reported by Pegler (1997) from the state of São Paulo, are also
excluded from the list of P. cubensis from Brazil, including M. Capelari MC248, which is too
insufficient for study so could not be confirmed, and G. Guzmán et al. 22947, which is
P. albostipitatus (Menolli et al. 2013). The collection of P. cubensis studied by Rosa et al.
(2003) and Rosa & Capelari (2009), although not listed in their works, was located at SP and
represents a species of sect. Hispidoderma, so far unidentified, with a trichodermal to
hymenidermal pileipellis of cystidioid elements, not a cutis as observed in P. cubensis.
Finally, a material deposited at JPB (I.C. Oliveira 286) as P. cubensis from the state of
Capítulo III - 99
Paraíba (northeast Brazil) actually represents a member of Entolomataceae (probably a
Rhodocybe sp.) and the material recorded by Xavier-Santos et al. (2004) as P. cf. cubensis is
not available at SJRP.
Recent molecular analyses (Justo et al. 2011a, b) including a Brazilian collection of
P. cubensis (CP233, as Pluteus sp. V – SP394389) showed its interesting position in sect.
Celluloderma like other taxa with cutis-like pileipellis that were previously assigned to sect.
Hispidoderma, viz. P. ephebeus (Fr.) Gillet and P. riberaltensis var. conquistensis Singer.
Pluteus eludens E.F. Malysheva, Minnis & Justo, Mycologia 103: 650. 2011.
Fig. 10
PILEUS 10–28 mm diam, convex when young, becoming applanate, sometimes subumbonate,
when fresh translucently striate < ½ R, black (5YR2.5/1) when very young and then fading to
dark greyish brown (10YR3/2) or dark brown (10YR3/3) at the centre and brown (7.5YR5/4)
to strong brown (7.5YR5/6), greyish brown/brownish orange (6CD3) or dark yellowish brown
(10YR4/4) elsewhere, surface smooth when young, becoming strongly areolate-venose at
centre with the veins rather distant and not reaching the pileus margin, margin not sulcate or
pectinate. LAMELLAE crowded, free, < 4 mm broad, at first white, finally greyish orange
(6B3), concolourous edges. STIPE 10–33(–45) × 1–3 mm, central, cylindrical to slightly
attenuated upwards, solid, greyish brown (5E3) or greyish white (2.5Y7.5/2) under densely
white translucently striate and pruinose-pubescent surface, dry. ODOUR weak. TASTE,
COLOUR
FLESH
and SPORE PRINT COLOUR not recorded. BASIDIOSPORES [40/2/2] 5.6–6.2(–7.5) × 5.0–
6.2 µm [Q = 1.00–1.24(–1.34); Qm = 1.10; Lm = 6.1 µm; Wm = 5.5 µm], globose to broadly
ellipsoid, rarely ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 21–26
× 10–15 µm, ventricose, thin-walled, four-spored. PLEUROCYSTIDIA 36–65 × 13.7–22 µm,
mostly lageniform or slightly clavate, thin-walled, colourless and hyaline, numerous.
CHEILOCYSTIDIA 22–50 × 8.7–15 µm, clavate, utriform or lageniform, thin-walled, colourless
or sometimes filled with pale brownish intracellular pigment, often with an apical mucilage,
crowded. PILEIPELLIS a hymeniderm composed of one layer of clavate to spheropedunculate
cells (25–47 × 12.5–24 µm) intermixed with elongate-fusiform, elongate-clavate or broadly
lageniform to filiform cystidioid elements (43–68 × 7.5–15 µm) with rounded apex and
evenly dissolved brown intracellular pigment. CAULOCYSTIDIA 27–36 × 11.2–13.7 µm, shortclavate to narrowly lageniform, filled with brownish-grey intracellular pigment. CLAMP
CONNECTIONS
absent.
Capítulo III - 100
HABIT, HABITAT AND DISTRIBUTION: Gregarious or in groups of two. In Paraná it was found in
mixed ombrophilous alluvial and montane forests, on decayed trunks of dicotyledonous, from
800 to 1,000 m above sea level. Fruiting in January (this record), although similar forms were
observed in the field from December to April by the second author, and between November
and January in the northern hemisphere (Justo et al. 2011c). Until now it was known from the
original description from Portugal, Russia and the U.S.A. (Justo et al. 2011c). Type locality:
Portugal, Madeira Island.
Fig. 10. Pluteus eludens (SJf-3912). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis elements.
e. caulocystidia. Bars = 10 µm.
Capítulo III - 101
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 28 Jan 1980, A.A.R. de
Meijer CUa-346 (MBM); Roça Velha, 6 Jan 2001, A.A.R. de Meijer SJf-3912 (MBM).
COMMENTS: Pluteus eludens is characterized by the combination of a pileipellis composed of
heteromorphic elements (sect. Celluloderma subsect. Mixtini according to Singer 1956, 1959,
1986) and pigmented cheilocystidia. Among the representatives of sect. Celluloderma, few
species combine these characters, including P. eugraptus and P. multiformis (Singer 1956,
1958, Pegler 1977, 1986, Justo et al. 2011c), but those two differ as follows: P. eugraptus has
non-globose basidiospores, a distinctly dark lamellar edge and cheilocystidia that are always
pigmented, and in P. multiformis pleurocystidia are scarce, cheilocystidia are mostly clavate
and the shape of the pileipellis cells is highly variable.
Although the presence of a dark brown lamellar edge and pigmented cheilocystidia are
mentioned in the protologue of P. eludens (Justo et al. 2011c), the constancy of these
characters is discussed by the authors, who emphasized that in specimens from the U.S.A. and
Russia have also been observed basidiomata with a concolourous or indistinctly pigmented
lamellar edge, whilst the cheilocystidia are mainly hyaline in collections from the U.S.A.
Thus, the presence of a concolourous lamellar edge and hyaline cheilocystidia should not
prevent us to identify our materials as P. eludens. Probably the most significant
morphological difference between the Brazilian collections and the materials studied by Justo
et al. (2011c) are the shape and size of the caulocystidia; they were described as
preponderantly cylindrical and lageniform (50–90 × 7.0–15.0 µm) in the holotype, whilst in
our collections they are short-clavate to narrowly lageniform (27–36 × 11.2–13.7 µm).
However, the taxonomic significance of this feature is unclear because caulocystidia were not
found in specimens of P. eludens from the U.S.A. and Russia (Justo et al. 2011c).
The Brazilian collections studied herein fit within the broad concept of P. eludens and it is the
first record of the species from South America.
Pluteus fibulatus Singer, Lilloa 25: 252. 1952.
Fig. 11
PILEUS 22–60 mm diam, at first slightly convex, then applanate, hygrophanous, when fresh
translucently striate up to the centre, centre and striae dark brown [10YR3/(3–4)], between
striae yellowish brown (10YR5/5), when dry centre dark brown (10YR3/3) and elsewhere
(very) pale brow/light yellowish brown (10YR6.5/3.5); surface smooth and dry. LAMELLAE
moderately close, free, < 9 mm broad, pinkish. STIPE 43–75 × 3–5 mm, central, very
Capítulo III - 102
gradually attenuated upwards, solid, white, dark brown innately fibrillose all over; surface
smooth and dry. FLESH white, < 3 mm thick near stipe and < 1 mm thick above midpoint of
lamellae. ODOUR indistinct. TASTE and
SPORE PRINT COLOUR
[60/3/2] 6.2–8.7(–9.3) × (5.0–)5.6–7.5(–8.1) µm [Q = 1.00–1.24(–1.40); Qm = 1.14; Lm = 7.4
µm; Wm = 6.5 µm], globose to broadly ellipsoid, occasionally ellipsoid, inamyloid, hyaline,
smooth, thick-walled, guttulate. BASIDIA 23–26 × 8.0–10.0 µm, versiform to clavate or
ventricose, thin-walled, four-spored. PLEUROCYSTIDIA 60–88 × 17.5–28.7 µm, fusoidventricose, metuloidal with thickened wall and apices provided with two to four apical
prongs, sometimes without prongs and with an acute or rounded apex, colourless and hyaline,
rather numerous. CHEILOCYSTIDIA 44–75 × 8.7–15.0 µm, clavate with rather long pedicel,
thin-walled, colourless, hyaline, abundant and mostly in fascicules. LAMELLAR
EDGE
sterile.
LAMELLAR TRAMA inverse, up to 50 µm wide, composed of thin-walled, hyaline hyphae, 2.5–
13.7 µm diam. PILEUS
CONTEXT
undifferentiated, up to 190 µm thick, composed of thin-
walled, hyaline hyphae, usually 3.7–13.7 µm diam or sometimes inflated up to 31 µm diam.
PILEIPELLIS a cutis up to 125 µm thick, composed of parallel chains of cylindrical, thin-walled
hyphae, individual terminal elements 30–60 × 7.5–20 µm diam, with rounded apex and evenly
dissolved brown intracellular pigment. CLAMP CONNECTIONS present.
HABIT,
Solitary to scattered. In Paraná it was found in mixed
ombrophilous alluvial forest, on decayed branches and stumps of dicotyledonous and trunks
of the palm Syagrus romanzoffiana (Cham.) Glassman, from 800 to 900 m above sea level,
fruiting in December, although similar forms were observed in the field until April. It is
known from Argentina (Singer and Digilio 1952, Singer 1959, Raithelhuber 1991) and Brazil
(Pegler 1997, Meijer 2006). Type locality: Argentina, Prov. Tucumán.
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 14 Dec 1979, A.A.R.
de Meijer CUa-284 (MBM); 26 Dec 1979, A.A.R. de Meijer CUa-284c (MBM).
COMMENTS: Pluteus fibulatus belongs to sect. Pluteus. It is apparently restricted to South
America and characterized by the fibrillose deeply coloured pileus and the presence of clamp
connections. During the re-examination of the collections from Paraná the first author could
not recover the clamp connections in the exsiccatae, but they were observed in fresh materials
by the second author. The presence of globose basidiospores observed in the collections from
Paraná could be a distinctive additional character; although Singer & Digilio (1952) and
Singer (1959) did not specify the basidiospores shape for P. fibulatus. A recent reexamination of the type collection of P. fibulatus (Menolli & Capelari 2013) demonstrated
that it contains globose to broadly ellipsoid or rarely ellipsoid basidiospores [20/1/1] (5.6–)
Capítulo III - 103
6.2–7.5(–8.7) × (5.0–)5.6–6.8(–7.5) μm [Q = 1.00–1.24(–1.40); Qm = 1.16; Lm = 7.1 μm;
Wm = 6.1 μm] like those observed in our materials. Menolli & Capelari (2013) found some
pleurocystidia of the Magnus-type in the type collection, which were not observed in the
collections from Paraná.
Fig. 11. Pluteus fibulatus (CUa-284). a. basidiospores. b. pleurocystidia. c. cheilocystidia. Bars = 10 µm.
Pluteus fibulatus seems to be a good species, although few collections have been identified
with certainty under this name and some morphological variation has been observed.
Undoubtedly, sect. Pluteus includes some species complexes and different morphological
interpretations have suggested surprising synonymies (Justo et al. 2011a, b, Menolli et al.
2013). For P. fibulatus probably only a study including molecular support will certify its
identity and its relationship with other species.
Capítulo III - 104
Pluteus fusconigricans (Berk. & Broome) Sacc., Syll. Fung. 5: 673. 1887.
Fig. 12
≡ Agaricus fusconigricans Berk. & Broome, Journ. Linn. Soc., Bot. 11: 534. 1871.
PILEUS 35 mm diam, concave, when fresh translucently striate < ½ R, centre dark brown
(6F4), elsewhere brown (6E4) to light yellowish brown (10YR6/4), extreme centre minutely
rugose-venose, elsewhere smooth, margin very slightly sulcate, dry. LAMELLAE close, free,
< 6 mm broad, at first pure white, soon pink; edge mostly entirely brown. STIPE 36 ×
3(middle)–4.5(base) mm, attenuated upwards, fistulose, base pale greyish brown, elsewhere
whitish, surface smooth, dry, glabrous. ODOUR,
COLOUR
TASTE, FLESH COLOUR
and
SPORE PRINT
not recorded. BASIDIOSPORES [20/1/1] 5.6–7.5 × 5.0–5.6 µm [Q = (1.00–)1.11–1.24
(–1.34); Qm = 1.19; Lm = 6.3 µm; Wm = 5.3 µm], suglobose to broadly ellipsoid, rarely
globose or ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 25–26 × 9.0
µm, clavate, thin-walled, four-spored. PLEUROCYSTIDIA 37–57 × 12.5–20 µm, clavate to
lageniform, mostly with evenly dissolved brown content or sometimes condensed mainly in
the apical or basal region, usually with apical mucilage to which adhere masses of
basidiospores, thin-walled, numerous. CHEILOCYSTIDIA 27–53 × 11.2–17.5 µm, clavate to
lageniform or vesiculose, with evenly dissolved brown content or sometimes condensed
mainly in the apical or basal region, thin-walled, numerous. PILEIPELLIS a hymeniderm
composed of one layer of thin-walled and clavate, spheropedunculate or subglobose cells, 41–
52 × 25–40 µm, with a short pedicel (2.5–6.2 µm long) and condensed brown intracellular
alluvial forest, on decayed branch of dicotyledonous, from 800 to 900 m above sea level.
Fruiting in April (this record) and in May and June in the Afrotropical and Indomalayan
regions, respectively (Pegler 1977). Previously it had been known only from Tanzania and Sri
Lanka (Pegler 1977, 1986). Type locality: Sri Lanka, Kandy Distr.
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 18 Apr 1980, A.A.R.
de Meijer CUa-434 (MBM).
COMMENTS: The combination of distinctly dark lamellar edge, suglobose to broadly ellipsoid
basidiospores, pigmented pleuro- and cheilocystidia and a hymenidermal pileipellis without
cystidioid elements fits the concept of P. fusconigricans (Pegler 1977).
Among the species that combine a distinctly dark lamellar edge with this type of pileipellis
and pigmented pleuro- and cheilocystidia, P. fusconigricans is only comparable with
P. luctuosus Boud. Recently, Justo et al. (2011b) showed the molecular relationship between
Capítulo III - 105
specimens that fit with the morphological concept of P. luctuosus and P. phlebophorus
(Ditmar) P. Kumm., considering they as possible synonyms. Nevertheless, the pleurocystidia
observed in our material are like those reported for P. fusconigricans instead of those of
P. luctuosus and P. phlebophorus that are longer (up to 100 µm long) and more lageniform
with long narrow neck (Orton 1986).
This paper reports P. fusconigricans for the first time from South America. Pluteus
fusconigricans belongs to sect. Celluloderma.
Fig. 12. Pluteus fusconigricans (CUa-434). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis
Pluteus glaucotinctus E. Horak, Bull. Jard. Bot. Belg. 47: 88. 1977.
Fig. 13
Capítulo III - 106
PILEUS 23–50 mm diam, at first hemispherical, then convex to plano-convex, finally
applanate and subumbonate, hygrophanous, when fresh translucently striate up to centre,
when dry non-striate, when young and fresh greyish yellow (3.5C3), when mature and fresh
in centre dark green (30F4) to olive brown (4E4 or 5Y4/3), or white (coll. ANc-4214),
elsewhere light grey [5Y6/(1-2)] to white (2.5Y8/2), striae pale brown (10YR6/3) to
yellowish grey (3B2), between striae pale to white, centre sometimes minutely flocculose,
smooth elsewhere, dry. LAMELLAE very close to close, free and remote from stipe,
subventricose, < 7 mm broad, at first white then orange white (5A2). STIPE 33–60 × 2–5 mm,
central, almost cylindrical to slightly attenuated from base upwards, solid, white, base with
extremely weak greenish blue tinge, bu when bruised distinctly staining bluish green (greyish
turquoise – 24D3), surface smooth to finaly striate, basal mycelium short tomentose, pure
white. FLESH greyish olive just under pileus surface, white elsewhere, < 2.8 mm thick near
stipe and < 0.5 mm thick above midpoint of lamellae. ODOUR fungoid. TASTE and
PRINT COLOUR
SPORE
not recorded. BASIDIOSPORES [100/5/5] 6.2–8.7 × 5.0–6.2(–6.8) µm
[Q = (1.10–)1.21–1.50; Qm = 1.31; Lm = 7.1 µm; Wm = 5.5 µm], broadly ellipsoid to
ellipsoid, rarely subglobose, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 18–
30 × 7.0–9.5 µm, clavate to urniform, thin-walled, four-spored. PLEUROCYSTIDIA 52–95
(–110) × 13.0–30 µm, ventricose or clavate, sometimes lageniform or utriform, more often
with truncate to subcapitate apex, colourless, hyaline, thin-walled but sometimes slightly
thickened mainly at apex. CHEILOCYSTIDIA 40–70 × 7.5–13.7 µm, ventricose-lageniform,
clavate or mucronate, usually elongated with well-developed pedicel, apex rounded or usually
with a short and narrow papilla, rarely truncate, thin-walled but usually with some incrusting
parietal pigment at apex, colourless and hyaline or sometimes seemingly with some colourless
amorphous internal content, crowded and mostly in fascicules. PILEIPELLIS a cutis composed
of cylindrical, thin-walled hyphae, 5.0–18.0 µm diam, apex usually obtuse, mostly with
evenly dissolved brownish intracellular pigment. CLAMP
CONNECTIONS
present and mostly
abundant in the covering layers.
HABIT, HABITAT AND DISTRIBUTION: Solitary, in groups of two or subgregarious. In Paraná it
was found in dense and mixed ombrophilous alluvial to montane forests, on decayed branches
and stumps of dicotyledonous, from 100 to 900 m above sea level, fruiting from January to
May. Considering previous records from other countries (Menolli et al. 2013), it seems to
fruit from August to May. Apparently it has a pantropical and mediterranean distribution
being known from the Democratic Republic of Congo (Horak 1977; Horak & Heinemann
1978), Brazil (Justo et al. 2011a, b), India and Spain (Menolli et al. 2013). Type locality:
Democratic Republic of Congo, Panzi-Kivu.
Capítulo III - 107
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 26 Jan 1980, A.A.R. de
Meijer CUa-426 (MBM); 7 Apr 1981, A.A.R. de Meijer CUa-287b (MBM); Antonina, Reserva Natural do Rio
Cachoeira, 21 May 2003, A.A.R. de Meijer ANc-4214 (MBM).
ADDITIONAL COLLECTION EXAMINED: BRAZIL, Rio Grande do Sul: Santa Maria, Boca do Monte, Passo do
Tigre, 16 Feb 2003, V.G. Cortez 001/03 (SMDB as P. nigrolineatus).
Fig. 13. Pluteus glaucotinctus. a. basidiospores (ANc-4214). b. pleurocystidium (CUa-426). c. cheilocystidia
(ANc-4214). Bar = 10 µm.
COMMENTS: Based on morphological and molecular data, Menolli et al. (2013) considered the
existence of at least three different species within a broad morphological concept of
P. glaucotinctus that makes it difficult to establish distinguishable features to reliably separate
them. The collections from Paraná fit the concept of P. glaucotinctus sensu lato and some
characteristics herein observed are shared with those observed in collections from different
geographic areas. For instance, the presence of clamp connections and cheilocystidia with
some incrusting parietal pigment at the apex are characteristics only observed in specimens
from India (Menolli et al. 2013). On the other hand, the presence of blue-green pigments was
neither observed in the collections from Paraná nor in those from the state of São Paulo
(Brazil) and India, but it was verified in those from Spain, Africa (Horak 1977, Horak &
Heinemann 1978, Menolli et al. 2013) and those from state of Rio Grande do Sul reported as
P. nigrolineatus by Wartchow et al. (2006). Similar morphological variation seems to occur
also with regard to the pigmentation of the cheilocystidia, only verified in the holotype
(Menolli et al. 2013).
Until now, based on morphological data we maintain the collections from Paraná under this
name. However, more studies using multiple evidences and, probably, more genes in the
molecular analyses will be necessary to propose a formal subdivision of the P. glaucotinctus
complex.
Capítulo III - 108
Although P. glaucotinctus was traditionally classified in sect. Hispidoderma (Singer 1986),
recent molecular studies confirmed its position in sect. Pluteus (Menolli et al. 2010, 2013,
Justo et al. 2011a, b).
Pluteus harrisii Murrill, Mycologia 3: 277. 1911.
Fig. 14
= Pluteus puttemansii Menolli & Capelari, Mycologia 102: 701. 2010.
PILEUS 15–36(–55) mm diam, applanate with slightly convex margin, sometimes
subumbonate, strongly hygrophanous, when fresh strongly translucently striate  ½ R, centre
and striae dark brown (7F5), between striae greyish red (7B3), when dry centre dark brown
(7F4) and elsewhere greyish brown (6E3); surface smooth to innately fibrillose, centre
sometimes minutely rugulose but mostly smooth. LAMELLAE close, free, < 4.5 mm broad, pale
orange (6A3). STIPE 20–60 × 2–5 mm, central, cylindrical, or slightly attenuated from base to
middle and cylindrical above middle, solid, white, sometimes very pale brownish
translucently striate; surface smooth, dry, macroscopically glabrous; basal mycelium forming
a short but dense white tomentum. FLESH white, in the fresh pileus concolourous with the
surface and drying pure white, < 1.5 mm thick near stipe and < 0.3 mm thick above midpoint
of lamellae. ODOUR indistinct. TASTE and SPORE
PRINT
[180/9/9] (6.2–)7.5–8.7(–10.6) × (5.0–)6.2–7.5(–8.7) µm [Q = (1.08–)1.16–1.40(–1.50);
Qm = 1.22; Lm = 8.1 µm; Wm = 6.6 µm], broadly ellipsoid to ellipsoid, rarely subglobose,
inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 21–31 × 7.5–11.2 µm, versiform
to clavate or ventricose, thin-walled, four-spored, sometimes with small scattered guttules.
PLEUROCYSTIDIA (40–)52–85(–91) × 12.5–25 µm, fusoid-ventricose, metuloidal with wall up
to 6.2 µm thick in upper part or 1.2 µm thick all over, apices usually with two to five
versiform lateral prongs (up to 6.2 µm long), prongs rarely with a secondary bifurcation or
with several small finger-like protuberances, occasionally without prongs and with a rounded
or irregular apex or with up to eight suberect, acute or obtuse horns, colourless and hyaline or
more often with a colourless internal condensation, moderately numerous to abundant.
CHEILOCYSTIDIA (26–)30–60 × 8.7–18.0 µm, clavate to versiform-clavate or clavatevesiculose, usually with a moderately long pedicel, thin-walled, colourless, hyaline, abundant
and mostly in fascicules. LAMELLAR
EDGE
sterile. LAMELLAR
TRAMA
inverse, up to 62 µm
wide, composed of thin-walled, hyaline hyphae, 2.5–13.7 µm diam, sometimes branched or
with terminal elements slightly inflated up to 17.5 µm diam. PILEUS
CONTEXT
undifferentiated, up to 200 µm thick, composed of thin-walled, hyaline hyphae, usually 3.7–
Capítulo III - 109
13.7 µm diam, sometimes inflated up to 31 µm diam. PILEIPELLIS a cutis up to 150 µm thick,
composed of parallel chains of cylindrical, thin-walled hyphae, individual terminal elements
40–70 × 12–20 µm, with rounded to sometimes attenuated obtuse apex (3.7–6.2 µm diam),
sometimes strongly constricted at the septa, with evenly dissolved brown intracellular
Fig. 14. Pluteus harrisii (MAg-3717). a. basidiospores. b. pleurocystidia. c. cheilocystidia. Bar = 10 µm.
HABIT, HABITAT AND DISTRIBUTION: Solitary to scattered. In Paraná it was observed in dense
and mixed ombrophilous forests, on decayed branches and trunks of dicotyledonous, at 800 m
above sea level. In Brazil its occurrence has been documented between November and April.
Tropical distribution: Jamaica and Cuba (Murrill 1911), Trinidad (Baker & Dale 1951 as
P. cervinus var. bambusinus), U.S.A. – Florida (Singer 1956, 1959), Guadeloupe (Pegler
1983), Mexico (Vargas et al. 1993; Rodríguez and Guzmán-Dávalos 2001) and Brazil (Pegler
1997; Meijer 2006 as P. aff. harrisii; Menolli et al. 2010). Type locality: Jamaica.
Capítulo III - 110
MATERIAL EXAMINED: BRAZIL, Paraná: Morretes, Parque Estadual Pico do Marumbi, Caminho do Itupava, 13
Jan 2000, A.A.R. de Meijer MAg-3717 (MBM).
ADDITIONAL COLLECTIONS EXAMINED: BRAZIL, Minas Gerais: Marliéria, Parque Estadual do Rio Doce, 16 Mar
2000, L.H. Rosa & R.O. Morais LHR102 (SP); São Paulo: São Miguel Arcanjo, Parque Estadual Carlos Botelho,
25 Apr 1986, M. Capelari & V.L.R. Bononi MC10 (SP); Santo André, Reserva Biológica de Paranapiacaba, 11
Apr 1990, M. Capelari et al. MC3282 (SP); 22 Mar 2007, Menolli Jr. et al. NMJ122 (SP); 24 May 2007, Menolli
Jr. et al. NMJ132 (SP); 06 Nov 2010, J.J.S. Oliveira & A.V. Costa JJSO267 (SP); São Paulo city, Parque
Estadual da Cantareira, Núcleo Engordador, 24 Apr 2007, Menolli Jr. et al. NMJ131 (SP – Holotype of
P. puttemansii); 31 Jan 2008, F. Karstedt et al. FK1066 (SP).
COMMENTS: Pluteus harrisii belongs to sect. Pluteus. It has been commonly recorded from
tropical areas and is characterized by moderately large basidiospores and metuloidal
pleurocystidia with poorly developed apical prongs (Banerjee & Sundberg 1995, Pegler 1983,
1997). However, recent molecular analyses (Justo et al. 2011b) related sequences from
specimens with this morphological concept to P. puttemansii Menolli & Capelari, which was
described with well-developed hooks in the metuloids. Although P. harrisii is apparently
considered a very variable species regarding the morphology of the pleurocystidia (Justo et al.
2011b), sequences from more collections that fit these morphological features will be
necessary to better define the concept of P. harrisii.
After the re-examination of collections identified as P. harrisii from Paraná, and also from
other Brazilian States, some corrections and considerations must be made. The collection
ANc-4219 previously mentioned by Meijer (2006) as P. cf. harrisii actually represents
P. albostipitatus. The collection L.H. Rosa & R.O. Morais PERD 090 studied by Rosa &
Capelari (2009) is in bad conditions of preservation and consequently insufficient to confirm
its identification, however it was possible to confirm the occurrence of P. harrisii in the state
of Minas Gerais based on the re-examination of the other material studied by them (LHR102).
Finally, the undetermined material (NMJ132) reported by Menolli et al. (2010) can be
considered P. harrisii.
Pluteus homolae Minnis & Sundb., N. Amer. Fung. 5: 37. 2010.
Fig. 15
≡ Prunulus ludovicianus Murrill, N. Amer. Fl. 9: 330. 1916.
≡ Mycena ludoviciana (Murrill) Murrill, Mycologia 8: 220. 1916.
PILEUS 15–18 mm diam, slightly convex and umbonate to applanate, translucently striate up
to centre, centre and striae brown (6F4), elsewhere brown (6E4), surface smooth, dry.
LAMELLAE close to moderately close, free, 4 mm broad, at first white, then pink, narrow edge
brown. STIPE 22–25 × 1.5–2 mm, cylindrical, sometimes slightly clavate at base, entirely light
Capítulo III - 111
brownish grey/light yellowish brown (2.5Y6/3) in coll. CUa-439, entirely white in coll. JSb3259, surface smooth, dry, glabrous. FLESH white. ODOUR indistinct. TASTE,
FLESH COLOUR
and SPORE PRINT COLOUR not recorded. BASIDIOSPORES [40/2/2] (5.0–)5.6–6.8 × 5.0–6.2 µm
[Q = 1.00–1.24; Qm = 1.08; Lm = 5.9 µm; Wm = 5.5 µm], globose to broadly ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 25–34 × 7.0–8.0 µm, clavate to
ventricose, thin-walled, four-spored. PLEUROCYSTIDIA 31–56 × 10.0–26 µm, clavate to
lageniform, colourless and hyaline, thin-walled, numerous. CHEILOCYSTIDIA 29–50 × 11.0–22
µm, clavate to broadly lageniform, usually with fuscous brown dissolved content, thin-walled,
abundant. PILEIPELLIS a hymeniderm composed of one layer of thin-walled and clavate,
lageniform or subglobose cells, 22–50 × (8.7–)10.0–21 µm, with a short pedicel (up to 8.7 µm
long) and condensed or more often evenly dissolved brownish intracellular pigment. CLAMP
CONNECTIONS
HABIT,
absent.
Solitary or in groups of two. In Paraná it was found in
mixed ombrophilous alluvial and seasonal semi-deciduous montane forests, on decayed
branches of dicotyledonous, from 600 to 900 m above sea level. Fruiting in February and
April (this record) and in September by the type collection (Minnis & Sundberg 2010). Until
now it was known only from the U.S.A. (Minnis & Sundberg 2010). Type locality: U.S.A.,
Louisiana.
MATERIAL EXAMINED: BRAZIL, Paraná: Jundiaí do Sul, Fazenda Pau d’Alho, 6 Feb 1996, A.A.R. de Meijer
JSb-3259 (MBM); São José dos Pinhais, Reserva Natural Cambuí, 20 Apr 1980, A.A.R. de Meijer CUa-439
(MBM).
COMMENTS: The globose to broadly ellipsoid basidiospores, pigmented cheilocystidia and
hymenidermal pileipellis without cystidioid elements fit the characteristics described for
P. homolae (Minnis & Sundberg 2010).
Pluteus submarginatus E. Horak described from Chile on decaying wood of Nothofagus sp.
also shares the combination of uniformly dark pigmented lamellar edges, colourless
pleurocystidia and a pileipellis without cystidioid elements (Horak 1964), but it is
distinguished by the more elongate basidiospores (7.2–8.5 × 5.6–6.8 µm) and the larger (70
–90 × 20–28 µm) and more lageniform pleurocystidia with long necks (Horak 1964).
This is the first record of P. homolae from South America and also the first record since its
description. Pluteus homolae belongs to sect. Celluloderma.
Capítulo III - 112
Fig. 15. Pluteus homolae (CUa-439). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis elements.
Bars = 10 µm.
Pluteus hylaeicola Singer, Fieldiana, Bot. 21: 95. 1989.
Fig. 16
PILEUS 62 mm diam, applanate and subumbonate, not hygrophanous, not translucently striate,
centre evenly brownish grey (6F2) and slightly rugose, outside centre innately fibrillose on
slightly paler brownish grey (6D2) background. LAMELLAE very close, free and 2 mm remote
from stipe, ventricose, < 7 mm broad, light brown (5D3), with very conspicuously brown
edge (visible without lens) that is concolourous with pileus centre. STIPE 60 × 5(apex)–
7(base) mm, central, base bulbous, above gradually attenuated to apex, solid, brownish grey
(6D2) innate striate on brown (5E4) background, surface dry, basal mycelium forming a white
tomentum. FLESH greyish white when fresh and white when dry, < 7 mm thick near stipe and
Capítulo III - 113
< 2 mm thick above midpoint of lamellae. ODOUR pleasant, possibly raphanoid. TASTE and
SPORE PRINT COLOUR not recorded. BASIDIOSPORES [20/1/1] 6.2–7.5 × (5.0–)5.6–6.2 µm [Q =
1.00–1.24; Qm = 1.12; Lm = 6.5 µm; Wm = 5.8 µm], globose, subglobose or broadly
ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA 25–27 × 9.0 µm,
clavate, thin-walled, four-spored. PLEUROCYSTIDIA (56–)61–81(–102) × (12.5–)17.5–25.7 μm,
fusoid-ventricose, metuloidal with wall mostly 1.2 µm thick all over, rarely < 4 µm thick at
apex, apices usually provided with two to five apical or lateral prongs up to 6.2 µm long, also
frequently clavate with rounded apex and quite thin-walled, particularly those near the
lamellar edge, colourless and hyaline, very abundant. CHEILOCYSTIDIA 49–58 × 12.5–17.5
µm, clavate to vesiculose, thin-walled, filled with brownish intracellular pigment, abundant
and mostly in fascicules. LAMELLAR
EDGE
sterile. LAMELLAR
TRAMA
inverse, up to 62 µm
wide, composed of thin-walled, hyaline, hyphae, 1.2–12.5 µm diam. PILEUS
CONTEXT
undifferentiated, up to 190 µm thick, composed of thin-walled, hyaline hyphae, up to 13.7 µm
diam. PILEIPELLIS a cutis composed of cylindrical, thin-walled hyphae, 5.0–11.2 µm diam,
with rounded apex and evenly dissolved brownish intracellular pigment. CAULOCYSTIDIA
absent. CLAMP
CONNECTIONS
present at all septa in the pileipellis and also abundant in the
hymenium.
HABIT, HABITAT AND DISTRIBUTION: Solitary. In Paraná it was found in dense ombrophilous
lowland forest, on a decayed dicotyledonous trunk, at 100 m above sea level. Fruiting in
August (this record) and in July (type collection, Singer 1989). It is known only from its
original description from Brazil (Singer 1989). Type locality: Brazil, Pará.
MATERIAL EXAMINED: BRAZIL, Paraná: Antonina, Reserva Natural do Rio Cachoeira, 10 Aug 2003, A.A.R. de
Meijer ANc-4248 (MBM).
COMMENTS: Pluteus hylaeicola is a species of sect. Pluteus characterized by pleurocystidia
without or with up to five sterigmatoid apical prongs, dimorphic and pigmented
cheilocystidia, absence of caulocystida and presence of clamp connections (Singer 1989). The
collection herein studied is slightly different from the type mainly with regard to the
cheilocystidia, which Singer (1989) described as being of two types: “(1) strongly elongate,
41.5–64 × 9.5–18 µm, subfusiform; (2) shortly ventricose-clavate, 23–74 × 11–36 µm,
generally 23–30 × 11–15 µm” (our translation). However, in our opinion the significance of
this feature is questionable because it can be considered a gradual variation within so similar
cheilocystidium shapes. Unfortunately, neither the holotype was located at BAFC, nor the
additional collection mentioned in the protologue was found at INPA.
Capítulo III - 114
Fig. 16. Pluteus hylaeicola (ANc-4248). a. basidiospores. b. pleurocystidia. c. cheilocystidia. Bars = 10 µm.
Among the species of sect. Pluteus with pigmented cheilocystidia, P. hylaeicola is
morphologically close to P. aporpus Singer, P. atromarginatus (Singer) Kühner,
P. atropungens A.H. Sm. & Bartelli, P. brunneoolivaceus E. Horak, P. eucryphiae,
P. martinicensis Singer & Fiard, P. raphaniodorus E. Horak, P. spegazzinianus Singer,
P. squamosopunctus E. Horak, P. subspinulosus E. Horak.
Although P. aporpus has pigmented cheilocystida, it has no clamp connections (Singer 1954a,
1959). Singer (1962) described P. aporpus f. porpophorus Singer that differs from the type
form in the presence of clamp connections, but it has smaller basidiospores (5.5–6 × 3.8–4.2
µm) than our material. Singer (1969) also suggested a probable relationship between
P. aporpus f. porpophorus and P. raphaniodorus, however the latter differes from
P. hylaeicola in having smaller basidiospores, 4.8–5.6 × 4–4.6 µm (Horak 1964). Pluteus
bruneoolivaceus has no clamp connections and its pileus is entirely covered by pyramidal
squamules (Horak 1964).
Capítulo III - 115
Pluteus atromarginatus has horned pleurocystidia only and usually grows on coniferous wood
(Singer 1956, Orton 1986, Banerjee & Sundberg 1995). Also P. squamosopunctus has horned
pleurocystidia only, which can be filled with brownish pigment (Horak 1964). Horak (1964)
described two additional forms of P. squamosopunctus, viz. P. squamosopunctus
f. alcaliodorus E. Horak and P. squamosopunctus f. magnocystis E. Horak, which also have
pigmented pleurocystidia. Another species with brownish cheilocystidia but exclusively
horned pleurocystidia is P. subspinulosus, which may even have lateral spinules (Horak
1964). Pluteus atropungens has pleurocystidia that are mostly unhooked with a rounded or
obtuse apex and rarely with one to three very short apical hooks (Justo & Castro 2007).
Pluteus martinicensis has fasciculate tufts of caulocystidia over the entire stipe surface
(Pegler 1983, Pradeep et al. 1996, 2002, Rodríguez & Guzmán-Dávalos 2007).
Pluteus spegazzinianus and P. eucryphiae are probably the species most closely related to
P. hylaeicola. Pluteus spegazzinianus has more elongate basidiospores (Q = approx. 1.23
–1.32) and longer pleurocystidia, 55–110 × 6.8–21 µm, mostly over 96 µm long (Singer 1952,
Singer & Digilio 1952). In addition, the brownish lamellar edge of P. spegazzinianus is
noticeable only after the investigation of several specimens with the aid of a hand lens,
because the cheilocystidia can also be hyaline (Singer 1952, Singer & Digilio 1952). Based on
ITS sequences, Justo et al. (2011b) recently suggested the relationship between
P. spegazzinianus and P. brunneidiscus Murrill / P. pouzarianus Singer, species characterized
by a pale lamellar edge combined with pigmented cheilocystidia. The main differences
between P. eucryphiae and P. hylaeicola are the occurrence in the former of pleurocystidia
with up to 10 spiniform prongs, incrusted pileipellis elements and cheilocystidia of one shape
(Singer 1969). Re-examining the holotype of P. eucryphiae [Chile, Valdivia, Cordillera
Pelada, R. Singer M5567, 7 May 1965 (BAFC)] we found subglobose to ellipsoid or rarely
globose basidiospores [20/1/1] (6.2–)6.8–8.7(–9.3) × 5.6–7.5(–8.1) µm [Q = (1.00–)1.07
–1.34(–1.40); Qm = 1.23; Lm = 7.9 µm; Wm = 6.5 µm] and metuloidal pleurocystidia mostly
without prongs or provided with 2–4(–6) apical or lateral prongs (Fig. 17). We could not
confirm the presence of incrusted pileipellis elements through the type re-examination and
few cheilocystidia were observed, as subhyaline elements filled with very pale yellow
dissolved pigment. Considering that P. eucryphiae is only known from the original
description and that some incongruities about its morphology could be noted, the real
relationship with P. hylaeicola will remain uncertain until more collections with these
characteristics have become available.
Capítulo III - 116
Although the specimens cited by Singer (1989) in the description of P. hylaeicola are lost, we
think the characteristics observed in the collection from Paraná are sufficient to identify it
with this name. We prefer not to assign the status of neotype for the Paraná collection until a
new topotypical material from the Amazonas region can be recovered.
Fig. 17. Pluteus eucryphiae (R. Singer M5567 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia.
Bars = 10 µm.
Pluteus neochrysaegis Menolli & de Meijer sp. nov.
Fig. 18
HOLOTYPUS: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 21 Apr 1980, A.A.R. de Meijer
CUa-442 (MBM).
ETYMOLOGY: The name refers to the morphological relationship with P. chrysaegis and to its occurrence in the
Neotropics.
PILEUS 15–30 mm diam, applanate, dry, not translucently striate, distinctly bicolourous, centre
dark brown/dark yellowish brown (10YR3/3.5), elsewhere brown/dark brown (10YR4/3),
centre minutely rugose-venose, elsewhere smooth, surface splitting radially and showing the
white flesh. LAMELLAE crowded, free to slightly remote, ventricose, 3 mm broad, white then
becoming pink. STIPE 15–50 × 3.5(middle)–5(base) mm, central, slightly attenuated from base
towards the apex, solid, surface innately fibrillose on white background, dry and smooth.
FLESH white. ODOUR indistinct. TASTE and
SPORE
PRINT
COLOUR
not recorded.
BASIDIOSPORES [40/2/2] (5.6–)6.2–7.5 × 4.3–5.6 µm [Q = (1.21–)1.24–1.36(–1.44); Qm =
1.28; Lm = 6.4 µm; Wm = 5.0 µm], broadly ellipsoid to ellipsoid, inamyloid, hyaline, smooth,
thick-walled, guttulate. BASIDIA 16–22 × 7.0–9.0 µm, clavate, four-spored. PLEUROCYSTIDIA
Capítulo III - 117
30–44 × 6.2–11.2 µm, versiform to lageniform or slightly ventricose, sometimes strangulated,
with rounded apex and usually subcapitate, with evenly dissolved light brown intracellular
pigment, slightly thick-walled like metuloids, wall up to 0.7 μm thick, very rare.
CHEILOCYSTIDIA dimorphic: I) 40–54 × 15–24 µm, clavate to lageniform or slightly
ventricose, usually with rounded apex, sometimes narrowly utriform, with pale yellow or light
brown
dissolved
intracellular
pigment,
thin-walled,
moderately
numerous;
II) 50–87 × 10–18.7 µm, fusiform, usually with very acute apex, with pale yellow or light
brown dissolved intracellular pigment, slightly thick- to thick-walled like metuloids of
Magnus-type, wall up to 1.2 μm thick, crowded. LAMELLAR EDGE sterile. LAMELLAR TRAMA
inverse, approx. 20 µm wide, composed of thin-walled, hyaline hyphae, up to 7.5 µm diam.
PILEIPELLIS a hymeniderm composed of thin-walled and clavate to vesiculose, or narrowly
utriform cells, 38–73 × 16.2–22 µm, with broadly rounded apex, mostly with evenly dissolved
brownish intracellular pigment but some also condensed. CAULOCYSTIDIA 25–47 × 10.0–16.2
µm, short-clavate to clavate, colourless or filled with pale yellow dissolved intracellular
HABITAT, HABIT AND DISTRIBUTION: Solitary or in groups of two. Only known from Paraná,
where it was found in mixed ombrophilous alluvial forest, on decayed wood of
dicotyledonous, at 870 m above sea level, fruiting in April and May.
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 21 Apr 1980, A.A.R.
de Meijer CUa-442 (MBM – Holotype); 27 May 1980, A.A.R. de Meijer CUa-442b (MBM).
COMMENTS: Pluteus neochrysaegis is characterized by a dark brown to dark yellowish brown
pileus, broadly ellipsoid to ellipsoid basidiospores, pigmented pleurocystidia that are rare and
slightly thick-walled, and pigmented cheilocystidia of two types differing in shape and wall
thickness (fusiform and thick-walled or clavate to lageniform and thin-walled).
The structure of the pileipellis observed in P. neochrysaegis places it in sect. Hispidoderma
and enforces its comparison with P. chrysaegis, another species of sect. Hispidoderma with
thick-walled cystidia (Pradeep et al. 2012). Different from P. neochrysaegis, P. chrysaegis
has a golden yellow pileus, brownish to yellowish stipe, globose to subglobose basidiospores,
colourless and shorter pleurocystidia (38–84 × 10.0–22.5 µm) that are abundant and evenly
distributed, and non-dimorphic cheilocystidia that are fusiform and colourless (Pegler 1986,
Pradeed & Vrinda 2006, Pradeep et al. 2012).
Capítulo III - 118
Fig. 18. Pluteus neochrysaegis (CUa-442 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia type I.
d. cheilocystidia type II. e. pileipellis elements. f. caulocystidia. Bars = 10 µm.
Pradeep et al. (2012) considered P. conizatus var. africanus E. Horak a synonym of
P. chrysaegis based on morphological and molecular data. Horak & Heinemann (1978)
separated P. conizatus var. africanus from the type variety mainly based upon differences in
the shape of the pileipellis elements and the pleurocystidia. P. conizatus (Berk. & Broome)
Sacc. var. conizatus also differs from P. neochrysaegis in having a yellowish brown stipe,
narrowly fusiform to lageniform pleurocystidia with up to 3 µm thick wall and cheilocystidia
Capítulo III - 119
similar to the pleurocystidia (Singer 1956, Pegler 1986). Moreover, Pegler (1986) described
the pileipellis of P. conizatus var. conizatus as “a disrupted trichodermium of narrow semierect to erect, septate hyphae, with terminal elements 20–42 × 4–7.5 µm”, clearly different
from the structure observed in P. neochrysaegis.
Pluteus rimosellus Singer, Lilloa 25: 262. 1952.
Fig. 19
= Pluteus subfibrillosus Singer, Trans. Brit. Mycol. Soc. 39: 187, 1956.
COMPLETE DESCRIPTION: Singer & Digilio (1952), Singer (1956, 1959).
MICROMORPHOLOGICAL
DATA FROM
PARANÁ STATE
COLLECTIONS:
BASIDIOSPORES [40/2/2]
6.2–7.5(–8.1) × 5.0–5.6 µm [Q = 1.21–1.50; Qm = 1.26; Lm = 6.4 µm; Wm = 5.1 µm],
broadly to ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA not
recovered. PLEUROCYSTIDIA (37–)45–65 × (6.2–)10.0–22 µm, ventricose-ampullaceous to
utriform, rarely clavate or strangulated, colourless and hyaline or usually with an internal
colourless to pale yellow intracellular condensation, thin-walled, abundant. CHEILOCYSTIDIA
25–52 × (8.7–)10.0–25 µm, clavate or slightly ventricose, mostly colourless and hyaline but
some with an internal colourless to pale yellow intracellular condensations, thin-walled,
abundant. PILEIPELLIS a trichoderm becoming eventually depressed like a cutis composed of
fascicles of cystidioid, ventricose-ampullaceous to broadly utriform, fusiform or clavate to
short-clavate terminal members, (30–)47–88(–106) × (11.2–)15.0–26 µm, with rounded to
sometimes attenuated obtuse apex, with evenly dissolved or sometimes condensed brownish
intracellular pigment. CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: Solitary to loosely gregarious. In Paraná it was found in
dense and mixed ombrophilous alluvial and lowland forests, on decayed dicotyledonous
trunk, from 100 to 900 m above sea level, fruiting in April and May. Singer & Digilio (1952)
and Singer (1959) reported it from September to January. Neotropical distribution: Argentina
(Singer & Digilio 1952, Singer 1959, Raithelhuber 1991) and Brazil (Singer 1956, 1959 as
P. subfibrillosus). Type locality: Argentina, Prov. Tucumán.
Meijer ANc-4215 (MBM); São José dos Pinhais, Reserva Natural Cambuí, 1 Apr 1980, A.A.R. de Meijer CUa417 (MBM).
Capítulo III - 120
Fig. 19. Pluteus rimosellus (CUa-417). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis
ADDITIONAL COLLECTIONS EXAMINED: ARGENTINA, Prov. Tucumán: Rio de los Sosas, 1 Jan 1951, R. Singer
T1089 (MICH, SP – Isotype of P. rimosellus); BRAZIL, Rio de Janeiro: Angra dos Reis, 30 Sept 1952,
R. Singer B433 (F – Holotype of P. subfibrillosus).
COMMENTS: Pluteus rimosellus belongs to sect. Hispidoderma and is characterized by a
fibrillose-rimulose pileus with sulcate margin, lamellar edge concolourous with the sides
(Singer & Digilio 1952, Singer 1959), globose to ellipsoid basidiospores, pigmented and
Capítulo III - 121
ventricose-ampullaceous to utriform or subcapitate pleurocystidia, clavate cheilocystidia and
a pileipellis composed of fascicules of cystidioid, ventricose-ampullaceous, fusiform or
clavate to short-clavate terminal elements.
Among the tropical species of sect. Hispidoderma with pigmented cystidia and lamellar edge
concolourous with the sides, P. rimosellus is to be compared with P. compressipes Murrill,
P. espeletiae Singer and P. subfibrillosus. The identity of P. compressipes is confusing
because some characteristics described in the protologue (Murril 1917) differ from those
observed during the re-examination of the type made by Singer (1956) and Smith & Stuntz
(1958), and also because the latter authors reported the presence of clamp connections,
although them to be rare. It is possible that P. rimosellus is a synonym of P. compressipes, but
as long as a new topotypical material remains unavailable, we prefer accept the name
P. rimosellus for the collections herein studied. Pluteus espeletiae differs from P. rimosellus
in having slightly larger basidiospores (8.0–8.8 × 6.8–7.5 µm) and a non-sulcate pileus
(Singer 1962).
Singer (1956, 1959) mentioned the closeness of P. subfibrillosus, P. rimosellus and
P. fibrillosus Murrill, but without mentioning how to distinguish them. Singer (1956) reported
the occurrence of pigmented cystidia in P. fibrillosus, but Smith & Stuntz (1958) and
Banerjee & Sundberg (1993) mentioned lack of internal pigment in the pleuro- and
cheilocystidia of the holotype. Besides, Smith & Stuntz (1958) and Banerjee & Sundberg
(1993) reported the occurrence of incrusted pileipellis elements in the holotype of
P. fibrillosus. These both characters were also verified in our re-examination of the type of
P. fibrillosus (see discussion under P. cubensis) and for us they seem sufficient to consider
P. fibrillosus as distinct from P. rimosellus and P. subfibrillosus.
With respect to P. rimosellus and P. subfibrillosus, most of the differences presented by
Singer (1959) are regard habitat and the macroscopic appearance, which may be considerable
characters in Pluteus (Justo et al. 2011b). Pluteus rimosellus was described with a cinereous
or brownish cinereous pileus, 10–35 mm diam, a pure white stipe and fruiting on Urticaceae,
while P. subfibrillosus was described with a porphyry brown, fuliginous, umber or dark grey
pileus, 30–60 mm diam, a stipe that is white in very young specimens but later becomes finely
brownish punctulate-fibrillose and the occurrence on dicotyledonous, but probably also on
Araucaria (Singer 1959). Our re-examination of the type collections of both species showed
that they share the same micromorphological characters, so we prefer treat them as synonyms.
Capítulo III - 122
The type of P. rimosellus (Fig. 20) has subglobose to broadly ellipsoid basidiospores [20/1/1]
(5.6–)6.2–7.5 × 5.0–6.2 µm [Q = 1.11–1.24(–1.30); Qm = 1.18; Lm = 6.4 µm; Wm = 5.4 µm],
pigmented and ventricose-lageniform to utriform pleurocystidia 27–50 × 7.5–13.7 µm, clavate
to slightly ventricose cheilocystidia 32–46 × 12.5–18.7 µm, and a pileipellis composed of
ventricose-utriform elements 67–72 × 15.0 µm, mostly with rounded apex and evenly
dissolved brownish intracellular pigment. The holotype of P. subfibrillosus (Fig. 21) has
broadly ellipsoid to ellipsoid basidiospores [20/1/1] 6.2–6.8 × 5.0–5.6 µm (Q = 1.21–1.36);
Qm = 1.26; Lm = 6.4 µm; Wm = 5.1 µm), pigmented and ventricose to utriform or rarely
clavate to strangulated pleurocystidia 43–56 × 13.7–18.7 µm, clavate to slightly utriform
cheilocystidia 29–41 × 11.2–20 µm, and a pileipellis composed of ventricose-utriform
elements 43–51 × 10.0–17.5 µm, mostly with rounded apex and evenly dissolved brownish
intracellular pigment.
Fig. 20. Pluteus rimosellus (R. Singer T1089 – isotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia.
d. pileipellis elements. Bar = 10 µm.
Based on the Paraná collections studied and on the synonymization proposed, this study
confirms the occurrence of P. rimosellus in Brazil. In the past, Singer (1954b) had identified
as P. rimosellus a sterile Brazilian collection (R. Singer B96) that was afterwards (Singer
1956) renamed as P. subfibrillosus. This record of P. rimosellus from Paraná corrects the
previous one (Meijer 2006) of collections CUa-417 and ANc-4215 misidentified as
P. cubensis.
Capítulo III - 123
Fig. 21. Pluteus subfibrillosus (R. Singer B433 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia.
d. pileipellis elements. Bar = 10 µm.
Pluteus riograndensis Singer, Lilloa 26: 116. 1954.
Fig. 22–23
= Pluteus beniensis Singer, Lloydia 21: 285, 1959.
PILEUS 4–65 mm diam, first ovoid with strongly involute margin touching the stipe, then
campanulate or slightly conic to plano-convex, eventually slightly depressed at centre,
concave or subumbonate, rugose-venose at first only in centre, soon all over surface and
radially arranged towards the margin, centre dark brown (6F6), margin (dark) brown (6EF4)
or sometimes paler, margin slightly sulcate or splitting radially and showing the white flesh,
sometimes apparently neither sulcate nor striate. LAMELLAE crowded to moderately distant,
free to sinuate-remote, ventricose, < 6 mm broad, first white then becoming orange
white/orange grey (6AB2), edge concolourous or sometimes distinctly dark only in few
points. STIPE 8–50 × 1–5(apex)–3–6(base) mm, central, slightly attenuated towards apex,
sometimes equal or with a small bulb, solid, sometimes becoming partially fistulose, when
young blackish brown at base and white elsewhere, when mature densely black brown
punctate towards base, or entirely white to translucently yellowish white striate, with scanty
Capítulo III - 124
basal mycelium. FLESH bluish grey, when dry white, < 1 mm tick near stipe and < 0.3 mm
thick above midpoint of lamellae. ODOUR indistinct. TASTE and
recorded.
BASIDIOSPORES
[320/16/16]
(4.3–)5.6–6.2(–7.5)
SPORE PRINT COLOUR
×
3.7–6.2(–7.5)
not
µm
[Q = 1.00–1.44(–1.68); Qm = 1.18; Lm = 6.1 µm; Wm = 4.9 µm), globose, subglobose to
ellipsoid, rarely elongate, inamyloid, hyaline, smooth, thick-walled. BASIDIA 18.7–37 × 6.2–
8.7(–10.0) µm, clavate to versiform, thin-walled, four-spored. PLEUROCYSTIDIA (32–)41–80
(–95) × (8.7–)10.0–29(–35) µm, clavate or ventricose to lageniform or fusiform to lanceolate,
colourless, sometimes filled pale yellowish intracellular content mainly those near the
lamellar edge, thin-walled, moderately numerous. CHEILOCYSTIDIA (20–)25–66 × (7.5–)10.0–
29(–33) µm, clavate to vesiculose, ventricose or slightly fusiform with a rounded apex,
utriform-pedunculate or subglobose to spheropedunculate, sometimes with basal septa,
colourless or frequently with light brown to dark-brown internal dissolved content, thinwalled, numerous. LAMELLAR EDGE heteromorphous. LAMELLAR TRAMA inverse, up to 62 µm
wide, composed of thin-walled, hyaline hyphae, 1.2–15.0 µm diam, sometimes with inflated
hyphae up to 20 µm diam. PILEUS CONTEXT undifferentiated, up to 125 µm thick, composed of
thin-walled, hyaline hyphae, 3.7–10.0 µm diam. PILEIPELLIS a hymeniderm up to 80 µm thick,
composed of one layer of thin-walled, spheropedunculate, clavate or subglobose elements,
25–63 × 12.5–49 µm, with a short to moderately long pedicel (2.5–15.0 µm long), mostly with
evenly dissolved brownish intracellular pigment but some also condensed. CAULOCYSTIDIA
34–47 × (8.7–)11.2–25 µm, clavate or subglobose, with evenly dissolved or sometimes
condensed brownish intracellular pigment, in few numbers. CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: Solitary to gregarious. In Paraná very common, in mixed
and dense ombrophilous alluvial and montane forests, on decayed branches, stumps and
trunks of dicotyledonous, from sea level to 950 m. From other Brazilian States fruiting from
October to April. Until now it was only known from the original description from Brazil
(Singer 1954b). Type locality: Brazil, Rio Grande do Sul.
MATERIAL EXAMINED: BRAZIL, Paraná: Curitiba, Bosque João Paulo II, 18 Mar 1992, A.A.R. de Meijer CUi2196; Guaraqueçaba, Tagaçaba, Reserva Natural Serra do Itaqui, 7 Nov 2009, A.A.R. de Meijer GUf-4480
(MBM); São José dos Pinhais, Associação Reserva Natural Cambuí, 3 Nov 1979, A.A.R. de Meijer CUa-208
(MBM); 20 Nov 1979, A.A.R. de Meijer CUa-208b (MBM); 21 Jan 1980, A.A.R. de Meijer CUa-208c (MBM);
22 Mar 1980, A.A.R. de Meijer CUa-405 (MBM);
ADDITIONAL COLLECTIONS EXAMINED: BRAZIL, Rio Grande do Sul: Passo Fundo, Jaboticabal, 16 Jan 2004,
M.S. Rother & B.M.A. Severo s.n (RSPF299 as P. beniensis); Santa Maria, Morro do Elefante, 2 Nov 2002,
D.L. Bordignon & J.C. Budke (SMDB9711 as P. beniensis); Santa Catarina: Blumenau, Parque Nacional da
Serra do Itajaí, Parque das Nascentes, Trilha da 3ª Vargem, 26 Jan 2011, F. Karstedt & R. Karstedt FK1800*
Capítulo III - 125
(SP); São Paulo: Canaeia, Parque Estadual da Ilha do Cardoso, 3 Feb 1987, D.N. Pegler et al. 3978 [K(M)];
Santo André, Reserva Biológica do Alto da Serra de Paranapiacaba, 14 Jan 2010, J.J.S. Oliveira & M. Capelari
JJSO09* (SP); 16 Mar 2010, J.J.S. Oliveira JJSO58* (SP); São Paulo city, Parque Estadual da Cantareira,
Núcleo Engordador, 12 Dec 2006, F. Karstedt & M. Capelari FK826* (SP as P. fuligineovenosus); 24 Apr 2007,
Menolli Jr. et al. NMJ127* (SP as P. fluminensis); 24 Apr 2007, Menolli Jr. et al. NMJ130* (SP as
P. jamaicensis); Parque Estadual das Fontes do Ipiranga, 24 Jan 2008, F. Karstedt & L.A.S. Ramos FK1046*
(SP as P. fluminensis).
Fig. 22. Pluteus riograndensis (GUf-4480). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis
elements. e. caulocystidia. Bars = 10 µm.
Capítulo III - 126
COMMENTS: Pluteus riograndensis belongs to sect. Celluloderma and is morphologically
close to other species described or recorded from South America by the presence of pleuroand/or cheilocystidia with fuscous content in all of them, such as P. beniensis, P. luctuosus
Boud. and P. rimosoaffinis.
Singer (1959) used the pigment condensation of the pileipellis elements as the principal
distinctive character to distinguish P. riograndensis and P. beniensis from Bolivia (the latter
described with evenly dissolved pigment), but after having re-examined a broad number of
collections we consider this character insufficient to distinguish these species. Moreover,
based on molecular data of ITS (data not show), we verified that sequence from specimens
(e.g. JJSO58) with pileipellis elements filled with evenly dissolved pigment are 99.83 %
identical to that from specimens (e.g. NMJ130) with condensed pigmentation (Fig. 23a, b).
Thus, we prefer to treat P. riograndensis and P. beniensis as synonyms. Pluteus beniensis was
described from Bolivia (Singer 1959) and since then only recorded from Brazil [Stijve & de
Meijer 1993, Meijer 2006 (in both as ‘P. cf. beniensis’), Wartchow et al. 2006]. The revision
of the collection mentioned in Stijve & de Meijer (1993) and Meijer (2006) was not possible
because this material was not located at MBM, but some of the Paraná collections herein
listed were previously published by Meijer (2006) as P. cf. fluminensis or as P. cf.
rimosoaffinis. For P. beniensis recorded by Wartchow et al. (2006), it was possible to observe
only basidiospores because the collection is poorly preserved and destroyed by insects.
After a careful re-examination of the collections previously identified as P. fluminensis Singer
by Pegler (1997) and as P. fluminensis, P. fuligineovenosus E. Horak and P. jamaicensis
Singer by Menolli et al. (2010), we concluded that almost of these materials fit the
morphological concept of P. riograndensis by the presence of pigmented queilocystidia,
although they are intermixed by some colourless. Only the collections MC164 and MC212
reported by Menolli et al. (2010) seem to be P. fluminensis, however.
Singer (1962) proposed P. riograndensis var. atromarginatus Singer from Argentina and
separated it from the type variety mainly by the black lamellar edge. All collection herein
studied have concolourous edge, but in some cases it was possible to see few points distinctly
dark on lamellar edge (Fig. 23c). New collections with entire dark edge and a comparative
study including molecular data will be necessary to certify the identity of P. riograndensis
var. atromarginatus and its relationship with other collections of P. riograndensis with
concolourous edge.
Capítulo III - 127
Fig. 23. Pluteus riograndensis. a. pileipellis elements with evenly dissolved pigment (JJSO58). b. pileipellis
elements with pigment condensation (NMJ130). c. lamellar edge with distinct darkly points. Bars a = 20 µm,
b = 100 µm.
Among the other species mentioned above, P. luctuosus differs from P. riograndensis in
having a distinctly coloured lamellar edge and slightly longer basidiospores (Singer & Digilio
1952, Singer 1959, Orton 1986, Homola 1972), but as discussed before, Justo et al. (2011b)
considered P. luctuosus and P. phlebophorus as possible synonyms. Pluteus luctuosus was
originally described from France (Boudier 1905) but also recorded from Argentina (Singer &
Digilio 1952, Singer 1956, 1959, 1962). Although, Singer (1962) mentioned that would be
necessary more collections to establish some relationship between the Argentinean material
and those from Europe. Pluteus rimosoaffinis also differs from P. riograndensis (among other
characters) by having the pileus surface cracking overall (Singer 1959).
Pluteus sapiicola Singer, Lloydia 21: 290. 1959.
Fig. 24
PILEUS 12–45 mm diam, plano-convex to applanate, slightly umbonate, rugulose at the centre
when fresh and becoming minutely wrinkled when dry, coffee brown to deep sepia brown
(N40Y70M40 to N60Y50M40) and slightly darker at margin (N40Y50M40 to N50Y80M80), margin
translucently sulcate-striate (< 2/3R) and sometimes finely cracking and showing the slightly
pinkish flesh. LAMELLAE free to almost free, moderately crowded, ventricose, < 3.5 mm
broad, white then becoming pale orange to pinkish (N10Y30M20), with one to three series of
lamellulae. STIPE 18–80 × 1.2–5(apex)–3–10(base) mm, central, sometimes curved,
cylindrical or slightly attenuated from base upwards, hollow, white and smooth at the apex,
sometimes brownish grey or greyish brown translucent striate on pale background, base
usually with greyish fibrils, slightly longitudinally striate, with strigose to tomentose pure
white basal mycelium. ODOUR fungoid. TASTE,
FLESH COLOUR
and
SPORE PRINT COLOUR
not
recorded. BASIDIOSPORES [140/7/7] 5.0–6.2(–8.1) × 5.0–6.2(–7.5) µm (Q = 1.00–1.12; Qm =
Capítulo III - 128
1.04; Lm = 5.8 µm; Wm = 5.6 µm), globose to subglobose, inamyloid, hyaline, smooth, thickwalled, guttulate. BASIDIA 20–42 × 6.0–8.7(–10.0) µm, clavate, thin-walled, four-spored.
PLEUROCYSTIDIA (37–)41–75(–83) × (13.7–)15.0–27 µm, lageniform to ampullaceous,
sometimes clavate or fusiform, as seen in KOH frequently with apical mucilage containing a
mass of basidiospores, thin-walled, colourless and hyaline, moderately numerous.
CHEILOCYSTIDIA (22–)27–56(–66) × (11.2–)15.0–20(–23) µm, like the pleurocystidia, clavate
to vesiculose, sometimes lageniform, ampullaceous or fusiform, thin-walled, colourless and
hyaline, not numerous. LAMELLAR
EDGE
apparently sterile with numerous basidioles.
LAMELLAR TRAMA inverse, up to 50 µm wide, composed of thin-walled, hyaline hyphae, 3.7
–6.2 µm diam. PILEUS
CONTEXT
undifferentiated, approx. 180 µm thick, composed of thin-
walled, hyaline hyphae, 2.5–6.2 µm diam, sometimes with oleiferous hyphae up to 5.0 µm
diam. PILEIPELLIS a hymeniderm up to 31 µm thick, composed of one layer of thin-walled,
spheropedunculate, clavate or subglobose elements, (27–)31–62 × (12.5–)17.5–42 µm, with a
short to moderately long pedicel (2.5–15.0 µm long), mostly with evenly dissolved brownish
intracellular pigment but some also condensed. CAULOCYSTIDIA 28–32 × 11.2–21 µm, clavate
or subglobose, with condensed brownish intracellular pigment, in few numbers (only
observed in M. Capelari s.n. PEFI52). CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: In Paraná it was found in mixed ombrophilous montane
forest, on decayed trunks of dicotyledons at 950 m above sea level. From the states of Paraná
and São Paulo fruiting between October and April. Until now it was only known from
Argentina (Singer 1959, Horak 1964, Raithelhuber 1991) and French Guyana (Courtecuisse
1991). Type locality: Argentina, Prov. Salta.
MATERIAL EXAMINED: BRAZIL, Paraná: Curitiba, Parque Barigui, 5 Mar 1993, A.A.R. de Meijer CUb-2552
(MBM); Colombo, Embrapa Florestas, 15 Jan 2001, A.A.R. de Meijer & M.A.L. de A. Amazonas COa-3929
(MBM).
ADDITIONAL COLLECTIONS EXAMINED: BRAZIL, São Paulo: São Paulo city, Parque Estadual da Cantareira,
Núcleo Engordador, 27 Apr 2006, F. Karstedt & M. Capelari FK629 (SP)*; 25 Oct 2009, M. Capelari & L.A.S.
Ramos MC4513 (SP)*; Parque Estadual das Fontes do Ipiranga, 20 Jan 1999, M. Capelari s.n. PEFI52 (SP);
Santo André, Reserva Biológica de Paranapiacaba, 21 Oct 2009, M. Capelari & L.A.S. Ramos MC4488 (SP)*;
8 Dec 2010, J.J.S. Oliveira et al. JJSO303 (SP)*.
COMMENTS: Pluteus sapiicola was described by Singer (1959) from xerophytic vegetation
and based on diagnostic characters including a striped stipe, pileipellis cells with evenly
dissolved pigment and ampullaceous cystidia usually with a cylindrical neck. Although Singer
(1959) considered the habitat as a distinctive character for P. sapiicola, since then this name
Capítulo III - 129
has also been applied to collections from wetter areas in Argentina (Horak 1964) and French
Guiana (Courtecuisse 1991).
Fig. 24. Pluteus sapiicola. a. basidiomata. b. basidiospores. c–d. pleurocystidia. e–f. cheilocystidia.
g–h. pileipellis elements. i. caulocystidia. a, f. MC4488. b, c, e, h. FK629. d, g. JJSO303. i. PEFI52.
Bars: a = 1 cm. b–i = 10 µm.
Pluteus sapiicola is morphologically close to P. pulverulentus Murrill and P. eliae Singer
with which it shares the mostly geometrically globose basidiospores. Courtecuisse (1991)
discussed the morphological relationship between these species and reported the presence of
Capítulo III - 130
caulocystidia in P. sapiicola that probably are responsible for the brownish grey colour of the
stipe. Dennis (1953) described a pigmented stipe and caulocystidia for P. pulverulentus from
Trinidad, but if stipe colour is a distinctive character for P. sapiicola, Dennis (1953) may have
seen P. sapiicola instead of P. pulverulentus, because the latter species is usually reported as
having a white stipe (Singer 1956, 1959, Pegler 1977, 1983). In spite of the brownish grey
stipe colour in almost all collections, we found caulocystidia only in the collection
M. Capelari s.n. PEFI52. A careful re-examination of the type material and more collections
with coloured stipe will be necessary to certify if the occurence of caulocystidia can be
considered a intraspecific variation in P. sapiicola.
According to Singer (1959), P. pulverulentus differs from P. sapiicola and P. eliae by the
presence of condensed pigmentation in the pileipellis elements. But the taxonomic
significance of this feature needs further studies because Pegler (1983), who also studied the
type, described pileipellis cells with evenly distributed content for P. pulverulentus. The
shape of the pleurocystidia would be different in P. pulverulentus and P. sapiicola: in the
former they are vesiculose, ventricose, lageniform or vesiculose-fusoid (Singer 1956, 1959,
Smith & Stuntz 1958, Pegler 1977, 1983) and in the latter they are ampullaceous with a
cylindrical neck (Singer 1959). Pluteus eliae is separated from P. sapiicola mainly by the
white stipe and shape of the cystidia.
Molecular evidences and more collections of Pluteus with these characteristics would be
required to establish the actual relationship between P. eliae, P. pulverulentus and
P. sapiicola. However, based on the stipe colour, pleurocystidium shape and the presence of
pileipellis cells with mostly evenly dissolved pigment, we prefer to maintain these collections
under P. sapiicola. Sequences from two collections herein studied (FK629 and MC4488)
were previously positioned (as Pluteus sp. II) in the sect. Celluloderma by Justo et al. (2011a,
b).
Pluteus striatocystis Pegler, Kew Bull. Addit. Ser. 6: 268. 1977.
Fig. 25
PILEUS 27 mm diam, convex, subumbonate, dark brown (5F3, 6F4), radially innate fibrillose
on white background, dry. LAMELLAE very close, free and 0.5 mm remote from stipe,
lanceolate, < 2.5 mm broad, at first white then becoming brownish orange (6C3). STIPE 45 × 3
mm, central, very slightly attenuated from base to apex, white, surface minutely appressed
white flocculose at lower half, striate, dry, basal mycelium pure white, downy. FLESH < 2mm
thick near stipe and < 0.2 mm thick above midpoint of lamellae. ODOUR, TASTE,
FLESH
COLOUR
Capítulo III - 131
and SPORE PRINT COLOUR not recorded. BASIDIOSPORES [20/1/1] 6.2–6.8 × 5.0–5.6 µm
[Q = (1.11–)1.21–1.24(–1.36); Qm = 1.23; Lm = 6.4 µm; Wm = 5.2 µm], broadly ellipsoid,
rarely subglobose or ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate.
PLEUROCYSTIDIA 40–70 × 15.0–25 µm, clavate or slightly ventricose to lageniform, colourless
and hyaline but some with an internal colourless to very pale yellow dissolved content,
frequently with a median or apical collar adhered to the wall and that appears longitudinally
striate in surface view, thin-walled, very abundant. CHEILOCYSTIDIA 36–46 × 11.2–20 µm,
clavate or slightly ventricose, colourless and hyaline but some with an internal colourless to
very pale yellow dissolved content, thin-walled, in few numbers. PILEIPELLIS a cutis
composed of parallel chains of cylindrical, thin-walled hyphae, individual terminal elements
up to 214 × 17.5 µm, usually with a subacute or obtuse apex (3.7 µm diam), mostly with
condensed brownish pigment but some also dissolved. CLAMP CONNECTIONS absent.
HABIT,
Solitary. In Paraná it was found in seasonal semi-
deciduous submontane forest, on litter, at 350 m above sea level. Fruiting in February (this
record) and in March and June in Africa (Pegler 1977). It was known only from the original
description from tropical Africa (Pegler 1977). Type locality: Kenya, Central Province.
MATERIAL EXAMINED: BRAZIL, Paraná: Fênix, Parque Estadual de Vila Rica do Espírito Santo, 24 Feb 1996,
A.A.R. de Meijer FE-3300 (MBM).
ADDITIONAL COLLECTIONS EXAMINED: KENYA, Central Province: Nairob District, City Park, 12 Mar 1968,
D.N. Pegler K6 [K(M) – Holotype].
COMMENTS: Pegler (1977) emphasized the longitudinally striate pleurocystidia as a distinctive
character to recognize P. striatocystis. Few differences were noted between the African and
the Brazilian collections: the pleurocystidia described from the holotype are smaller (26–46 ×
8.0–16 µm) than those from Paraná. In the latter material could be that the striae in the
pleurocystidia are due to an outer collar adhered to the wall. After slight pressure on
microscopic slides, the lamellar sections fragmented and many collars detached from the
pleurocystidia. Pegler (1977) thought that in P. striatocystis the striae are part of the true wall,
but the re-examination of the holotype showed that the striae are due to an outer collar,
exactly as in this material from Paraná. The type of P. striatocystis has subglobose to ellipsoid
basidiospores [20/1/1] 5.6–7.5(–8.7) × 4.3–6.2 µm [Q = 1.12–1.44; Qm = 1.27; Lm = 6.4 µm;
Wm = 5.1 µm], weakly pigmented and lageniform or rarely clavate pleurocystidia 50–69 ×
13.7–24 µm, and weakly pigmented and clavate or slightly ventricose cheilocystidia 39–50 ×
10.0–18.7 µm (Fig. 26).
Capítulo III - 132
Fig. 25. Pluteus striatocystis (FE-3300). a. basidiospores. b. pleurocystidia. c. cheilocystidia. d. pileipellis
elements. Bars: = 10 µm.
Pluteus diptychocystis is also characterized by an outer collar adhered to the wall (Singer
1954a, 1956, 1959, 1969). After a re-examination of the type collection of P. diptychocystis
[Argentina, Patagonia, Parque Nacional Nahuel Huapi, Isla Victoria, R. Singer M725, 17 May
1952, (MICH, SP – Isotype)], we verified that it can be distinguished from P. striatocystis
mainly by the shape of the basidiospores and the characteristics of the collar in the
pleurocystidia. The type of P. diptychocystis has broadly ellipsoid to ellipsoid basidiospores
[20/1/1] 6.8–8.7 × 5.0–6.2 µm (Q = 1.21–1.50; Qm = 1.33; Lm = 7.3 µm; Wm = 5.5 µm] and
Capítulo III - 133
most of the pleurocystidia have an outer collar, as illustrated by Singer (1956) that is never
striate.
Fig. 26. Pluteus striatocystis (D.N. Pegler K6 – holotype). a. basidiospores. b. pleurocystidia. c. cheilocystidia.
Bar: = 10 µm.
Recent molecular analyses (Justo et al. 2011a, b) led to some rearrangements to the traditional
infrageneric classification proposed by Singer (1959, 1986): as has been the case with
P. cubensis, P. ephebeus and P. riberaltensis var. conquistensis (see discussion under
P. cubensis), it is most likely that also P. striatocystis will turn out to be a member of sect.
Celluloderma. The Brazilian material herein studied was previously reported by Meijer
(2006) as P. cubensis. However, because of the characteristic pleurocystidia this material
clearly fits with the morphological concept of P. striatocystis. The present paper reports for
the first time this species from Brazil and represents its first record outside Africa.
Pluteus sublaevigatus (Singer) Menolli & Capelari, Mycology 1: 148. 2010.
≡ Pluteus chrysophlebius subsp. sublaevigatus Singer, Lloydia 21: 278. 1958.
ILLUSTRATION: Singer (1959), Menolli et al. (2010).
Capítulo III - 134
PILEUS 11–28 mm diam, campanulate then convex to plane, sometimes subumbonate or
usually concave at the centre, hygrophanous, when fresh strongly translucently striate > ½ R,
mainly on point of insertion of the lamellae, yellow elsewhere (N00A30M40 to N00A90M50),
centre strongly rugulose. LAMELLAE close, free to subfree, ventricose, < 3.5 mm broad,
yellowish, usually with one lamellula for each lamella. STIPE 28–33 × 1–4 mm, central,
cylindrical or slightly attenuated from base towards apex and with a subbulbous base,
yellowish white to light yellow (N00A90M50) with dark yellow base, surface smooth, minutely
striate and sometimes with white punctations and white pruinose mycelium at the base.
ODOUR indistinct. FLESH
COLOUR, TASTE
and
SPORE PRINT
[140/7/5] (5.0–)5.6–7.5 × (5.0–)5.6–7.5 µm [Q = 1.00–1.11(–1.24); Qm = 1.04; Lm = 6.3 µm;
Wm = 6.1 µm], globose to subglobose, rarely broadly ellipsoid, inamyloid, hyaline, smooth,
thick-walled, guttulate. BASIDIA (20–)22–27(–30) × 7.5–8.7(–10.0) µm, clavate, thin-walled,
four-spored. PLEUROCYSTIDIA 34–65 × 12.5–28.7 µm, clavate to vesiculose, sometimes
slightly ventricose, usually with an internal colourless to pale straw dissolved content, thinwalled, not numerous. CHEILOCYSTIDIA (22–)31–55(–62) × (7.5–)10.0–27(–32) µm, clavate to
vesiculose, sometimes slightly ventricose or subglobose, usually with an internal colourless to
pale straw dissolved content, thin-walled, numerous. LAMELLAR
TRAMA
EDGE
sterile. LAMELLAR
inverse, up to 62 µm wide, composed of thin-walled hyphae, 3.7–10 µm diam,
hyaline. PILEUS
CONTEXT
undifferentiated, approximately 187 µm thick, composed of thin-
walled hyphae, 2.5–6.2(–16.2) µm diam, hyaline. PILEIPELLIS a hymeniderm up to 50 µm
thick, composed of one layer of thin-walled and clavate, vesiculose, spheropedunculate or
subglobose cells, (25–)35–55 × (16.2–)21–34 µm, with a short pedicel (2.5–7.5 µm long),
colourless and hyaline. CLAMP CONNECTIONS absent.
HABIT, HABITAT AND DISTRIBUTION: Gregarious or in groups of two. In Paraná it was found in
dense ombrophilous submontane forest, on decayed trunk of dicotyledons of Astrocaryum
aculeatissimum (Schott) Burret (Arecaceae), at 300 m above sea level, fruiting in April.
Considering the type and other records from Brazil it occurs from January to April. Pluteus
sublaevigatus is known from Bolivia (Singer 1959) and Brazil (Menolli et al. 2010). Type
locality: Bolivia, Dpto La Paz.
MATERIAL EXAMINED: BRAZIL, Paraná: Morretes, BR-277, 15 Apr 1993, A.A.R. de Meijer Mae-2611 (MBM).
ADDITIONAL COLLECTIONS EXAMINED: BOLIVIA, Dpto. La Paz: Prov. Nor-Yungas, Charobamba, 13 Feb 1956,
R. Singer 1147 (LIL — Holotype). BRAZIL, Minas Gerais: Nova Ponte, Fazenda Caxuanã, Reserva Borda do
Rio Claro, 22 Jan 2007, R. Maziero s.n. (SP416737); São Paulo: São Paulo city, Parque Estadual da Cantareira,
Núcleo Engordador, 19 Feb 2008, F. Karstedt et al. FK1085 (SP)*; Parque Estadual das Fontes do Ipiranga,
1 Mar 2011, F. Karstedt & J.J.S. Oliveira FK1904 (SP)*.
Capítulo III - 135
COMMENTS: Re-examining all Brazilian collections, and differently from what was previously
described by Menolli et al. (2010), we verified the occurrence of cystidia that are slightly
coloured like those observed in P. globiger Singer.
Current molecular data from species with yellow basidiomata from Brazil (data not shown),
revel the existence of at least four putative species in this complex, including the already
described P. aureovenatus Menolli & Capelari, P. globiger and P. sublaevigatus. Thus, for
now, we consider P. sublaevigatus a good species characterized mainly by the presence of a
yellow pileus without brown or red-orange tints, predominantly globose basidiospores,
slightly coloured cystidia and colourless pilleipelis cells.
Pluteus thomsonii (Berk. & Broome) Dennis, Trans. Brit. Mycol. Soc. 31, 206. 1948. Fig. 27
≡ Agaricus thomsonii Berk. & Broome, Ann. Mag. Nat. Hist. 17: 131. 1876.
PILEUS 14 mm diam, applanate and subumbonate, dry, translucently striate at the margin,
centre yellowish brown (5F4½), elsewhere brownish grey (5D2), non-striate when dry,
surface reticulately venose (fresh and dry), particularly so in centre but with veins reaching
outer margin. LAMELLAE very close, free, ventricose, < 3 mm broad, at first white, becoming
pink. STIPE 14 × 3(middle)–5(base) mm, central, cylindrical or attenuated from base towards
apex, base broadened, solid but becoming hollow, white to whitish and dark brown punctate
all over, surface dry and smooth, basal mycelium white tomentose. FLESH greyish brown.
ODOUR indistinct. TASTE and
SPORE PRINT COLOUR
not recorded. BASIDIOSPORES [20/1/1]
6.2–7.5(–8.7) × 5.6–7.5 µm [Q = 1.00–1.11(–1.19); Qm = 1.05; Lm = 6.6 µm; Wm = 6.3 µm],
globose to subglobose, rarely broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled,
guttulate.
BASIDIA
30–40
×
8.0–11.0
µm,
ventricose-pedicellate,
four-spored.
PLEUROCYSTIDIA apparently absent. CHEILOCYSTIDIA (31–)55–76 × 8.7–20 µm (including
apical projection), clavate to slightly ventricose, usually with a long apical projection (up to
62 µm long), sometimes with an amorphous colourless content at apex, numerous. PILEIPELLIS
a hymeniderm composed of thin-walled and clavate, vesiculose, ventricose, fusiform or
subglobose cells, 46–88 × 15–26 µm (including apical projection), sometimes with a long
apical projection (up to 50 µm long) particularly those near the pileus centre, most elements
with pale yellow content condensed near the apex, but some colourless and hyaline.
CAULOCYSTIDIA not seen. CLAMP CONNECTIONS absent.
Capítulo III - 136
alluvial forest, on decayed trunks of dicotyledonous, from 800 to 900 m above sea level, with
fruiting only in December. The only other Brazilian record is from October (Wartchow et al.
2004). Of widespread distribution, having been reported from Africa (Vellinga 1990,
Breitenbach & Kränzlin 1995), Asia (Imazeki et al. 1988, 2011, Breitenbach & Kränzlin
1995, Yang et al. 2011), Europe (Orton 1960, 1986, Vellinga & Schreurs 1985, Vellinga
1990, Breitenbach & Kränzlin 1995), North America (Homola 1972, Minnis & Sundberg
2010) and South America (Wartchow et al. 2004). Type locality: England, West Kent.
MATERIAL EXAMINED: BRAZIL, Paraná: São José dos Pinhais, Reserva Natural Cambuí, 16 Dec 1979, A.A.R.
de Meijer CUa-292 (MBM).
COMMENTS: Pluteus thomsonii is characterized by a very dark and usually strongly reticulatevenose pileus, cheilocystidia with short to quite long apical projections, pleurocystidia also
usually with an apical projection but scarce to rare or apparently absent, pileipellis cells with
elongate-clavate to fusiform-ventricose elements that sometimes have a long apical projection
(Homola 1972, Orton 1986, Minnis & Sundberg 2010). The characteristics observed in our
collection fit the broad morphological concept of P. thomsonii except for the absence of
caulocystidia, which is a character that should be investigated for taxonomic importance.
Wartchow et al. (2004) reported P. thomsonii from Brazil based on a material in which
caulocystidia were also wanting. Recently, Rodríguez et al. (2008) described P. neotropicalis
O. Rodr.-Alcántar based on specimens from Mexico previously misidentified as P. thomsonii
(Rodríguez & Guzmán-Dávalos 1999) and the authors suggested that the specimen studied by
Wartchow et al. (2004) also should be considered P. neotropicalis instead of P. thomsonii.
According to Rodríguez et al. (2008) P. neotropicalis differs from P. thomsonii mainly in the
length of the projection of the cheilocystidia, which is shorter in P. neotropicalis (5.0–28 µm
long). Unfortunately, Wartchow’s collection was not available at SMDB for examination.
Molecular data of P. thomsonii are available only for specimens from the northern hemisphere
and recent analyses have confirmed its position in sect. Celluloderma (Justo et al. 2011a, b)
and showed that it involves a species complex composed of a number of secondary clades
within a single P. thomsonii morphological concept (Justo et al. 2012). Based solely on
morphological data and in spite of the absence of caulocystidia, for the time being we prefer
to keep this collection from Paraná as P. thomsonii. However, only molecular studies will
show the real relationship between the South American specimens and those from the
northern hemisphere.
Capítulo III - 137
Fig. 27. Pluteus thomsonii (CUa-292). a. basidiospores. b. cheilocystidia. c. pileipellis elements. Bars: = 10 µm.
Pluteus xylophilus (Speg.) Singer, Lilloa 22: 405. 1951.
Fig. 28
≡ Entoloma xylophilum Speg., Bol. Ac. Nac. Cord. 28: 305. 1926.
= Pluteus xylophilus var. tucumanensis (Singer) Singer, Lloydia 21: 209. 1959.
= Pluteus xylophilus var. major Singer, Lloydia 21: 210. 1959.
PILEUS 30–115 mm diam, convex to plano-convex, centre often subumbonate or umbonate,
becoming applanate or even slightly concave, not hygrophanous, dry, smooth to indistinctly
innate-fibrillose, extreme centre smooth to sometimes appressed flocculose, dark brown (7F4)
or sometimes fading to greyish yellow (4C4) and paler towards the margin, margin sometimes
inflexed, not sulcate or striate. LAMELLAE remote, crowded to close, subventricose,  16 mm
Capítulo III - 138
broad, at first white then becoming pinkish, with lamellulae up to four lengths. STIPE 23–150
× 3–13(apex)–9–20(base) mm, central, attenuated upwards, base sometimes strongly clavate
to subbulbous, solid, white to cream at apex, increasingly dense brown (7E4) appressed
fibrillose downwards, basal mycelium short tomentose and pure white. FLESH pure white,
 11 mm thick near stipe and  4 mm thick above midpoint of lamellae. ODOUR fungoid to
slightly raphanoid. TASTE mild. SPORE
PRINT
brownish orange (6C4). BASIDIOSPORES
[220/22/21] (5.0–)5.6–7.5(–8.7) × 3.7–5.0(–6.2) µm [Q = (1.21–)1.24–1.68(–1.74); Qm =
1.47; Lm = 6.5 µm; Wm = 4.5 µm], broadly ellipsoid to ellipsoid, occasionally elongate,
inamyloid, hyaline, smooth, thick-walled, guttulate. BASIDIA (18.7–)21–30 × 6.5–8.7 µm,
versiform to clavate, thin-walled, four-spored, with small scattered guttules and usually with
very short sterigmata. PLEUROCYSTIDIA 41–87 × (8.7–)10.0–25 µm, fusiform-ventricose or
lageniform to fusiform, metuloidal with wall 0.8–3.0 μm thick in upper part or entire wall
evenly thickened, colourless and hyaline, usually three types distinguishable regarding the
apex features: I) Cervinus-type with two to six apical or lateral prongs, abundant and
frequently with basidiospores adhered to apices; II) like those as found in P. harrisii, usually
without prongs and subcapitate or sometimes with two short lateral prongs, moderately
abundant; III) Magnus-type with an acute to obtuse apex, scarce to rare. CHEILOCYSTIDIA
21–115 × 7.0–15.0 µm, ventricose-utriform, cylindrical or clavate to subclavate, sometimes
with a small mucro at apex or with a long pedicel, thin-walled, colourless and hyaline, usually
crowded, in some collections scarce. LAMELLAR
EDGE
heteromorphous. LAMELLAR
TRAMA
inverse, up to 62 µm wide, composed of cylindrical and thin-walled, hyaline hyphae, 1.2–20
µm diam. PILEUS CONTEXT undifferentiated, thin to very thick up to 1,000 µm, composed of
thin-walled, hyaline hyphae, 2.5–21 µm diam. PILEIPELLIS a cutis up to 125 µm thick,
composed of parallel chains of cylindrical and thin-walled hyphae, individual terminal
elements 30–90 × 3.7–13.0 µm, with rounded to obtuse apex and brown intracellular pigment.
STIPITIPELLIS composed of strictly cylindrical, thin-walled hyphae, 3.0–5.0 µm diam. CLAMP
CONNECTIONS
HABIT,
absent.
HABITAT, AND DISTRIBUTION:
Solitary to scattered. In Paraná it occurs in mixed
ombrophilous montane and alluvial and dense ombrophilous submontane forests, on decayed
dicotyledonous stumps and trunks, sometimes on clayey soil near decayed dicotyledonous
wood, from sea level to 1,000 m. Fruiting has been documented and observed year round.
Neotropical distribution being previously recorded from Argentina (Spegazzini 1925 as
Entoloma xylophilum), Bolivia (Singer 1959), Brazil (Bresadola 1920 as P. cervinus var.
brasiliensis, Grandi et al. 1984, Stijve & de Meijer 1993, Pegler 1997, Meijer 2006, 2009,
Capítulo III - 139
Menolli et al. 2010), Mexico (Ramírez-Guillén & Guzmán 2003) and Peru (Singer 1959).
Type locality: Argentina, Prov. Buenos Aires.
Fig. 28. Pluteus xylophilus. a. basidiospores (CUa-160). b. pleurocystidia (Anc-4165). c. cheilocystidia
(SJf-4023). Bar: = 10 µm.
MATERIAL EXAMINED: BRAZIL, Paraná: Antonina, Reserva Natural do Rio Cachoeira, 20 Apr 2003, A.A.R. de
Meijer ANc-4165, (MBM); São José dos Pinhais, Reserva Natural Cambuí, 3 Oct 1979, A.A.R. de Meijer CUa160 (MBM); 16 Nov 1979, A.A.R. de Meijer CUa-160b (MBM); Roça Velha, 26 Feb 2001, A.A.R. de Meijer
SJf-4023 (MBM); 28 May 2001, A.A.R. de Meijer SJf-4086 (MBM).
ADDITIONAL COLLECTIONS EXAMINED: Rio Grande do Sul: Arroio do Meio, 1920, Rick s.n. (SP33926 as
P. cervinus); Passo Fundo, 16 Jan 2004, M.S. Rother & B.M.A. Severo s.n. (RSPF294); São Leopoldo, 1934,
Rick s.n. (PACA14516 as P. cervinus); 1906, Rick s.n. (PACA14519 as P. cervinus); 1934, leg. Steffen s.n.
(PACA14526 as P. cervinus); São Paulo: Mogi Guaçu, Reserva Biológica e Estação Experimental de Mogi
Guaçu, Gleba A, 9 Nov 2011, G.L. Robledo et al. s.n. (SP417741); Piracicaba, 15 Apr 1972, Milanez s.n.
(SP112156); São Paulo city, Parque Estadual das Fontes do Ipiranga, 5 Mar 1975, L.C. Abreu 283 (SP as
Volvariella bakeri); 10 Nov 1982, G. Guzmán 22986 (SP); 16 Jan 1987, Pegler et al. 3712 (SP); 20 Jun 2006,
F. Karstedt et al. FK683 (SP)*; 30 May 2007, Menolli Jr. et al. NMJ138 (SP)*; 18 Oct 2007, Menolli Jr. &
F. Karstedt NMJ150 (SP); Parque Estadual da Cantareira, Núcleo Engordador, 21 Aug 2007, Menolli Jr. et al.
NMJ143 (SP)*; 23 Oct 2008, M. Capelari & L.A.S. Ramos MC4397 (SP)*; 5 Jan 2012, M. Capelari & P.O.
Ventura MC4655 (SP)*.
COMMENTS: Pluteus xylophilus is a species of sect. Pluteus commonly recorded from South
America and characterized by a moderately large pileus, narrow basidiospores and three types
of pleurocystidia. Pluteus xylophilus used to be characterized also by the presence of
dimorphic cheilocystidia (Singer 1959, Menolli et al. 2010), but after having re-examined a
broad number of collections we consider the differences observed represent a gradual
Capítulo III - 140
variation within different shapes of cheilocystidia. Thus, the two varieties of P. xylophilus
proposed by Singer (1959), viz. P. xylophilus var. tucumanensis and P. xylophilus var. major,
are here considered synonyms also due to the extreme variation in size and pigmentation of
the pileus observed in our collections.
Recent morphological studies have shown that many Brazilian specimens previously
identified as P. cervinus actually are P. xylophilus (Menolli & Capelari 2013). This paper also
includes under P. xylophilus a specimen misidentified as Volvariella bakeri (Murrill) Shaffer
(Grandi et al. 1984, Pegler 1997) that later reported as Pluteus sp. by Menolli & Capelari
(2008).
Acknowledgements
The authors thank the curators of BAFC, F, JPB, K, LIL, NY, MBM, MICH, PACA, RSPF, SMDB for the loan
of collections studied; Carla Puccinelli, Fernanda Karstedt and Jadson José Souza de Oliveira for collecting
some specimens; Klei R. Sousa for preparing the illustrations; the Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) for the fellowships to N. Menolli Jr. and M. Capelari; and the Fundação de
Amparo à Pesquisa do Estado de São Paulo (FAPESP grant 2009/53272-2) for financial support.
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CAPÍTULO IV
Pluteus sect. Hispidoderma in Brazil
with new records based on morphological and molecular data*
Nelson Menolli Jr., Alfredo Justo & Marina Capelari
*Artigo a ser submetido para publicação na revista Cryptogamie Mycologie
Capítulo IV - 147
Capítulo IV - 148
Pluteus sect. Hispidoderma in Brazil
with new records based on morphological and molecular data
Nelson MENOLLI Jr. a,b, Alfredo JUSTO c & Marina CAPELARI b
a
CCT/Biologia, Rua Pedro Vicente 625, 01109-010, São Paulo, SP, Brazil; email:
[email protected] (corresponding author)
b
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902,
São Paulo, SP, Brazil; email: [email protected]
c
Clark University, Biology Department, 950 Main St., Worcester, 01610 MA, USA; email:
[email protected]
Running title: Pluteus sect. Hispidoderma in Brazil
Abstract – A revision of Pluteus sect. Hispidoderma occurring in Brazil is presented showing
the occurrence of eight species considered certainly known in the country: P. argentinensis,
P. chusqueae, P. longistriatus, P. maculosipes, P. neochrysaegis, P. rimosellus, P. varzeicola
and P. velutinus. Molecular analyses using ITS sequences showed the phylogenetic position
of P. argentinensis, P. cf. fernandezianus and P. velutinus, which are reported for the first
time from Brazil. Additionally, P. maculosipes also represent a new record based on
morphological data.
Key words: Agaricales / biodiversity / ITS / Pluteaceae / taxonomy
Capítulo IV - 149
INTRODUCTION
Singer (1959, 1986) considered the infrageneric classification of Pluteus with three
sections (Pluteus, Hispidoderma Fayod and Celluloderma Fayod) based on morphological
features such as the structure of the pileipellis and characteristics of the pleurocystidia. The
section Hispidoderma was recognized by Singer (1959, 1986) to include species with nonmetuloid pleurocystidia and pileipellis composed of elongated elements forming a cutis, a
hymeniderm or a trichoderm.
Vellinga & Schreurs (1985) proposed a variation of this classification subdividing
the sect. Hispidoderma into two taxonomic units according to the structure of the pileipellis.
A new section, viz. sect. Villosi Schreurs & Vellinga, was introduced to accommodate the
species with a cutis-like pileipellis, and the remaining species were transferred to sect.
Celluloderma as members of subsection Hispidodermini (Fayod) Vellinga & Schreurs that
includes species with a trichodermic pileipellis or a hymeniderm with cylindrical to fusiform
elements.
Singer’s classification is t e most accepted for infrageneric organization of Pluteus
and it has been supported by molecular data (Menolli et al., 2010; Justo et al., 2011a, b) with
some rearrangements. Justo et al. (2011a, b) showed in the molecular analyses that the species
with non-metuloid cystidia and a cutis-like pileipellis should be classified in sect.
Celluloderma instead of sect. Hispidoderma or sect. Villosi as proposed by Singer (1959,
1986) and Velllinga & Schreurs (1985), respectively. Thus, Justo et al. (2011a, b)
characterized the sect. Hispidoderma as having hymenidermal or trichodermal pileipellis
composed of long and elongated elements, which are very variable in shape and size.
Based on this circumscription, species of sect. Hispidoderma have been reported
from Brazil by Rick (1919, 1930, 1938, 1961), Singer (1954, 1956, 1959, 1989), Pegler
(1997), Meijer (2006), Wartchow et al. (2006), Rosa & Capelari (2009), Menolli & Capelari
(2010) and Menolli et al. (2013a, b). The purpose of this study is to improve the knowledge of
Pluteus sect. Hispidoderma in Brazil providing new records based on morphological and
molecular data and presenting a list of all species previously published in the literature with
an update of the taxonomic status of the Brazilian collections.
MATERIALS AND METHODS
The materials studied in the morphological examination include specimens recently
collected in Brazil or previously published. The herbarium acronyms follow Thiers (2013)
and t e ‘ ede
Capítulo IV - 150
rasileira de Herb rios’ (http://www.botanica.org.br/rede_ herbarios.php) for
the Herbarium of the Passo Fundo University (RSPF).
Species characterized as having non-metuloid pleurocystidia and a pileipellis as a
cutis were excluded because we accept the section delimitation as complemented by Justo et
al. (2011a, b) that considered these species in sect. Celluloderma. Some species with
pleurocystidia of thin- to slightly thick-walled and a cutis-like pileipellis were also not studied
here because they were considered members of sect. Pluteus, viz. P. albostipitatus (Dennis)
Singer, P. glaucotinctus E. Horak and P. nigrolineatus Murril (Menolli et al., 2010, 2013a;
Justo et al., 2011a, b).
The macroscopic description was based on fresh specimens. Colour terms are
according to Küppers (1979). In some cases the macroscopic description of the Brazilian
collections was not recovered but other references are provided below the taxonomic entry to
complement the species recognition. For microscopic analyses, the dried material was wetted
wit 70% et anol and t en re ydrated in 5% KOH or stained wit Melzer’s reagent to
determine the amiloidity reaction of the basidiospores. The notation [a/b/c] at the beginning of
a set of basidiospore data is to be read as “(a basidiospores were measured from (b)
basidiomata taken from (c collections”. Q represents t e range of t e lengt /widt ratio for
all of the measured spores, Qm represents the average of all calculated Q values for all of the
measured basidiospores and Lm (Wm) represents the average of all of the lengths (widths) of
the measured basidiospores. At least 20 basidiospores from each basidioma were measured in
lateral view, and the terms denoting basidiospore shape follow Bas (1969).
Methods for DNA isolation, PCR and sequencing follow Justo et al. (2011b). The
ITS region was amplified using the primer pair ITS1-F and ITS4 (White et al., 1990; Gardes
& Bruns, 1993). The molecular analyses were conducted with existing DNA sequences used
by Justo et al. (2011a, b), Pradeep et al. (2012) and from newly collected sequences of
materials representing species of sect. Hispidoderma. Newly obtained sequences were
deposited in GenBank. All GenBank accession or collection numbers and sequence
geographic origins are given in Fig. 1. Sequences were aligned using MAFFT version 6
(http://mafft.cbrc.jp/alignment/server/; Katoh & Toh, 2008) with the Q-INS-i option. Then,
the alignment was visually examined and manually corrected using MacClade 4.05
(Maddison & Maddison, 2002). Maximum Parsimony (MP) and Maximum Likelihood (ML)
were performed with the parameters specified in Justo et al. (2011b). Pluteus diettrichii Bres.
and P. seticeps (G.F. Atk.) Singer were used as outgroup taxa. Sequence divergence was
calculated using MatGAT (Campanella et al., 2003).
Capítulo IV - 151
RESULTS
The final dataset consists of 71 ingroup sequences with a total of 766 characters
(gaps included), of which 280 are parsimony informative. In the MP analysis 1792 MPT were
recovered (Length = 1027; CI = 0.57; RI = 0.90).
The current knowledge of sect. Hispidoderma in Brazil, as accepted in this paper, is
presented in Table 1 and the section ‘Ta onomy’ is reserved to include only the new records
from Brazil. The phylogenetic position of P. argentinensis, P. cf. fernandezianus,
P. longistriatus and P. velutinus is highlighted in Fig. 1 and discussed in detail for the new
records under Taxonomy. Additionally, P. maculosipes is reported for the first time from
Brazil based on morphological data.
The species identification of many non-Brazilian sequences presented in the tree
must be revised and some other yet to be identified, but it falls outside the scope of this paper.
Most of these sequences belongs to the leoninus/flavofuligineus/roseipes complex and others
are marked with “***” in Fig. 1.
Searches of the bibliographical and herbarium records led to a total of 19 taxa names
of sect. Hispidoderma that were linked to specimens collected in Brazil (Table 1). However,
only eight species are here considered certainly known in Brazil. The record of
P. cf. fernandezianus is maintained as uncertain determination due to the lack of macroscopic
data for the specimen studied and to few micromorphological differences verified between the
protologue (Singer, 1959) and our collection. Additionally, the record of P. aquosus needs
new collections to certify its existence in Brazil (Table 1). The occurrence of the other nine
taxa were not confirmed because the collections associated to these names were not located,
are too insufficient to study, or represent species of other sections. Previous studies (Menolli
& Capelari, 2013; Menolli et al., 2013a, b) re-examined some collections of sect.
Hispidoderma species, disregarding their occurrence in Brazil.
The infrageneric classification of some species should be regarded with caution. It is
not clear to us the real position of P. aquosus, P. polycystis and P. sergii but based on
characteristics of the pileipellis described by Singer (1956, 1959) we prefer to maintain them
in sect. Hispidoderma. Nevertheless, the occurrence P. polycystis and P. sergii could not be
confirmed in Brazil (Table 1). The same statement is applied to P. exiguus and its variety
because we consider the pileipellis structure as a palisade or a trichoderm (Vellinga and
Schreurs 1985, Orton 1986) to classify them in sect. Hispidoderma.
Capítulo IV - 152
Table 1: Species of Pluteus sect. Hispidoderma recorded from Brazil and notes about the identification of the
Brazilian collections.
Species
Reference
P. aquosus Singer a, b
P. argentinensis
P. chusqueae (E. Horak) Menolli c
P. exiguus (Pat.) Sacc. a
P. exiguus var. venosus Rick, nom. inv. a
Wartchow et al. (2006)
this paper
Menolli et al. (2013b)
Rick (1938, 1961)
Rick (1961)
P. cf. fernandezianus
P. granulatus Bres.
this paper
Rick (1930, 1938, 1961)
P. haywardii Singer
P. haywardii
P. cf. haywardii
P. leoninus
Pegler (1997)
Rosa & Capelari (2009)
Meijer (2006)
Rick (1938, 1961)
P. longistriatus (Peck) Peck c
P. maculosipes
Menolli & Capelari (2010),
Menolli et al. (2013a)
this paper
P. neochrysaegis Menolli & de Meijer c
P. cf. polycystis Singer a
Meijer (2006) d
P. rimosellus Singer c
P. sergii Singer a
Singer (1954), Menolli et al.
(2013b)
Pegler (1997)
P. subfibrillosus Singer
Singer (1956, 1959)
P. umbrosus (Pers.) P. Kumm.
Rick (1919, 1938, 1961)
P. varzeicola Singer
P. velutinus
Pluteus sp.
Singer (1989)
this paper
this paper (NMJ193) e
Notes about identification of
the Brazilian collections
too insufficient for study
not found
most likely P. jamaicensis
(Menolli & Capelari 2013)
see comments in Taxonomy part
most likely P. glaucotinctus E.
Horak (Menolli & Capelari,
2013)
not found
most likely P. conizatus (Berk. &
Broome) Sacc. (Menolli &
Capelari, 2013)
not found / too insufficient for
study (Menolli et al. 2013b)
P. albostipitatus (Menolli et al.,
2013a)
synonym of P. rimosellus
(Menolli et al., 2013b)
Singer (Menolli & Capelari,
2013)
Names in bold are considered species certainly known in Brazil. a See discussion below about the infrageneric
classification of these species.
b
Species that need new collections to certify their existence in Brazil. c Species
not studied in this work but that are considered as occurring in Brazil due to the well-preserved collections and
the complete description presented by the authors referenced. d Reference that includes two or more collections
that are actually different species. e Collection to insufficient for morphological identification, but included in the
current molecular analyses.
TAXONOMY
Pluteus argentinensis Singer, Lloydia 21: 230, 1959.
Figs. 2a–c, 3, 4
Pileus 30–52 mm diam, conic to plane-convex with a distinct umbo or almost
papillate, surface pale brown (N80A50M50) with darker centre (N90A99M50) and paler towards
Capítulo IV - 153
Fig. 1. Best tree from the maximum likelihood analysis for the ITS dataset of Pluteus sect. Hispidoderma.
Bootstrap values ≥70 % are s own on branc es (ML/MP). Root length has been reduced to facilitate graphical
representation. *** Voucher collections that need revision for correct identification.
Capítulo IV - 154
the margin, covered by appressed and radially arranged fibrils, sometimes exposing at the
margin the pale to translucent background between the fibrils, not translucently striate or only
so at extreme margin. Lamellae free, pinkish (N10A30M30), crowded, ventricose, with
concolourous edges and lamellulae of different lengths, 1–3 for each lamellae. Stipe 30–70 ×
2–3(apex)–5–7(base) mm, slightly tapering towards the apex, central, cylindrical, surface
white to pale cream with greyish to brownish base, longitudinally striate and sometimes with
brownish fibrils (N30A40M20) at the base, with or without scanty basal mycelium. Odour,
taste and flesh colour not recorded.
Basidiospores [60/3/3] 5.0–6.2 × (4.3–)5.0–5. μm (Q = 1.00–1.30; Qm = 1.20; Lm
= .0 μm; Wm = 5.0 μm , globose to broadly ellipsoid, inamyloid, hyaline, smooth, thickwalled, guttulate. Basidia 26–36 × 6.2–7.5 μm, elongate-clavate, thin-walled, four-spored.
Pleurocystidia 41– 7(−75 × 12.5−29 μm, ventricose to ampullaceous or rarely clavate,
colourless and hyaline, thin-walled, sparse and not abundant. Cheilocystidia 30–65 × (8.7–)
12.5–22(–27) μm, fusiform, slightly ventricose to slightly utriform or clavate, sometimes
subcapitate and with some incrusting parietal pigment at apex, colourless and hyaline, thinwalled, moderately abundant. Lamellar trama inverse, up to 31 μm wide, composed of thinwalled hyphae, 2.5–12.5 μm diam, interwovened by oleiferous hyphae up to 2.5 μm diam,
hyaline. Pileus context undifferentiated, appro . 50 μm t ick, composed of thin-walled
hyphae, 2.5–15.0 μm diam, yaline. Pileipellis a trichohymeniderm up to 125 μm t ick,
composed of fusoid or elongate-clavate cells, individual terminal elements (74–)104–162 ×
(10–)20–30 μm, with acute to subacute apex (up to 4.0 μm diam) or few with broadly rounded
apex (up to 10.0 μm diam), sometimes with points of apical incrustation, with evenly
dissolved or sometimes concentrated brown intracellular pigment. Caulocystidia 37–64(–81)
× 12.5–1 .2 μm, fusiform to ampullaceous or mucronate, with rounded or gradually
attenuated apex (2.5–6.2 µm diam), thin-walled, filled with dissolved brownish content.
Clamp connections absent in all parts examined.
Habit and habitat. – Subgregarious to disperse (up to three basidiomata near), on
decayed wood or on soil.
Revised specimens. – BRAZIL: Minas Gerais State, Marliéria city, Parque Estadual
do Rio Doce, 16 February 2000, L.H. Rosa & R.O. Marais PERD76 (SP307730); São Paulo
State, São Paulo city, Parque Estadual das Fontes do Ipiranga, 1 March 2011,
F. Karstedt & J.J.S. Oliveira FK1903 (SP417462); 3 March 2011, F. Karstedt & J.J.S.
Oliveira FK1915 (SP417457).
Remarks. – Pluteus argentinensis was described from Argentina (Singer, 1959) and
since then also reported from Mexico (Rodríguez & Guzmán-Dávalos, 1999, 2001). Wright &
Capítulo IV - 155
Albertó (2002) additionally reported it from Pampas in Argentina, but this record is doubtful
because they did not found pleurocystidia and illustrated the pileipellis elements and basidia
with clamp connections.
The protologue of P. argentinensis (Singer, 1959) includes few differences when
compared to the characteristics of the Brazilian collections, such as a slightly smaller pileus
(22–35 mm diam) and slightly bigger basidiospores (6.2–7.5 × 4.8– .5 μm , but certainly
without significance to certify its occurrence in Brazil.
Fig. 2. a–c. Pluteus argentinensis (FK1903); d–f. P. maculosipes (FK1651); g–i. P. velutinus (FK1889). Scale
bars: 1 cm.
Interestingly, the collection FK1915 was found growing on soil. Singer (1959)
suggested a humicolous habitat when he compared P. argentinensis to P. hispidulus
f. terrestris Kühner and by characterizing it in the key as growing on wood humus. Although
the difference in habitat, no morphological differences were observed between the other
Capítulo IV - 156
collections studied to consider the occurrence of a second taxa. Unfortunately, we were not
able to obtain ITS sequence from FK1915 and therefore P. argentinensis is maintained as
growing on soil and on wood until it is possible the molecular comparison between specimens
from different habitats.
Fig. 3. Pluteus argentinensis (FK1903). a. basidiospores; b. pleurocystidia; c. cheilocystidia. Scale bars: 10 µm.
According to Singer (1959), P. argentinensis is morphologically close to
P. fuliginosus Murrill, P. pluvialis and P. yungensis Singer. However, P. fuliginosus has some
pronged pleurocystidia (Smith & Stuntz, 1958; Banerjee & Sundberg, 1993), P. pluvialis has
a long plicate-sulcate pileus and pure white stipe (Singer, 1959), and P. yungensis has bigger
basidiospores [5.8–8.2 × 5.0–7.0(–7.5 μm] and w ite stipe w en fres (Singer, 1959 .
Phylogenetically, P. argentinensis is part of the species-complex round P. plautus
(Weinm.) Gillet but it is still unclear to which species it is more closely related (Fig. 1).
Capítulo IV - 157
Pluteus cf. fernandezianus Singer, Lloydia 21: 220, 1959.
Figs. 5, 6
Basidiospores [20/1/1] 6.2–8.7 × 5.0–7.5 μm (Q = 1.00–1.34; Qm = 1.18; Lm = 7.3
μm; Wm = .3 μm , globose to ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate.
Basidia 30–36 × 10–12.5 μm, clavate or versiform, thin-walled, four-spored. Pleurocystidia
52–104 × 15.0−35 μm, ventricose to lageniform or elongate-clavate to slightly fusiform,
sometimes with some incrusting parietal pigment at apex, usually filled with pale brownish
content, thin-walled, numerous. Cheilocystidia 44–54 × 18.7–25 μm, fusiform or utriform,
filled
with
pale
brownish
content,
thin-walled,
not
numerous.
Lamellar
edge
heteromorphous. Pileipellis a hymeniderm composed of one layer of versiform elements,
terminal cells 29–67 × 13.7–29 μm, vesiculose, clavate to fusoid cells (approx. 46–67 × 13.7
–29 µm) or in few number intermixed by spheropedunculate cells (approx. 29–38 × 24–29
µm), with rounded apex, thin-walled, mostly hyaline and colourless but some filled with pale
straw dissolved content. Stipitipellis a cutis of thin-walled hyphae, 2.5–8.7 µm wide,
colourless and mostly hyaline. Clamp connections absent in all parts examined.
Habit and habitat. – solitary on decayed wood.
Revised specimen. – BRAZIL: Rio Grande do Sul State, Passo Fundo city,
Jaboticabal, 13 September 2003, M.S. Rother & B.M.A. Severo s.n. (RSPF330).
Remarks. – The collection RSPF330 was deposited as P. beniensis Singer, a species
of sect. Celluloderma characterized by an epithelial pileipellis with uniform elements, but the
molecular analysis together with the morphological data confirms this mistake and its position
in sect. Hispidoderma. Pluteus cf. fernandezianus is also placed in the plautus clade (Fig. 1).
Despite the lack of macroscopic data for this collection, we tentatively related it to
P. fernandezianus described from Chile (Singer, 1959). Nevertheless, our collection has some
differences from the holotype on the micromorphological characteristics, such as the presence
of smaller pleurocystidia (46–65 × 8.7−23.3 μm), ellipsoid to short ellipsoid but non-globose
basidiospores [7–8(–9.3) × 6–6.7(–8) μm], and the lack of spheropedunculate cells in the
pileipellis as described for P. fernandezianus by Singer (1959). After, Singer (1969)
considered P. brunneoolivaceus E. Horak as synonym of P. fernandezianus but without any
discussion of the reasons for this. Horak (1964) described P. brunneoolivaceus from
Argentina and reported the presence of small horns and finger-like protuberances in the
pleurocystidia, which is probably similar to those described for P. fernandezianus by Singer
(1959 as aving “t e tip sometimes s ort apiculate in t e center, sometimes sinuate or
nodulose at ape ”. T e pleurocystidia of our collection usually ave apical incrustation
(Fig. 5b) but they are different from the apical ornamentation described and illustrated by
Horak (1964) for P. brunneoolivaceus. An accurate morphological comparison including
Capítulo IV - 158
revision of the type will be necessary to state the identity of P. cf. fernandezianus and to
establish the real relationship between P. brunneoolivaceus and P. fernandezianus.
By the presence of pigmented pleuro- and cheilocystidia in P. cf. fernandezianus, it
could be compared to other species of the stirps umbrosus (Singer, 1986) such as P. jaffuelii
(Speg.) Singer, P. umbrosus, P. multistriatus Murrill., P. fibrillosus Murrill, P. subfibrillosus,
P. compressipes Murrill and P. espeletiae Singer, but all of them also has morphological
differences that prevents the correct identification of P. cf. fernandezianus.
Fig. 5. Pluteus cf. fernandezianus (RSPF330). a. basidiospores; b. pleurocystidia; c. cheilocystidia. Scale bars:
10 µm.
Capítulo IV - 159
Fig. 6. Pileipellis elements of Pluteus cf. fernandezianus (RSPF330). Scale bar: 10 µm.
Pluteus jaffuelii (Singer, 1959) has smaller pleurocystidia [(30)–48–68 × 10−24 μm],
smaller basidiospores (6.2–7.5 × 4.8−7 μm) and much longer pileipellis elements (66–150 ×
13.8−30.8 μm). Pluteus umbrosus is a species commonly reported from Europe (Orton 1986)
with smaller basidiospores (5–7.5 × 4.5−6 μm) and much longer pileipellis elements (90–250
× 12−30 μm). Pluteus umbrosus and related species represent a separate lineage in sect.
Hispidoderma (Fig. 1). Pluteus multistriatus, alt oug
aving pleurocystidia up to 110 μm
long (Singer, 1956), was described by Singer (1956) and Smith & Stuntz (1958) as having
smaller basidiospores (6.5–7 × 4.8−5 μm and 5.4–6.5 × 4.7−5.5 μm, respectively) and with
hyaline cheilocystidia (Smith & Stuntz, 1958). Pluteus fibrillosus has smaller basidiospores
(5.8–7 × 5.5−6.8 μm, according to Smith & Stuntz, 1958), pleuro- and cheilocystidia lacking
internal pigment (Smith & Stuntz, 1958; Banerjee & Sundberg, 1993), and incrusted
Capítulo IV - 160
pileipellis elements (Banerjee & Sundberg, 1993; Menolli et al., 2013b). Pluteus
subfibrillosus was considered synonym of P. rimosellus by Menolli et al. (2013b) and
according to these authors the type has smaller basidiospores (6.2–6.8 × 5.0–5.6 µm). Pluteus
compressipes was described by Singer (1956) and Smith & Stuntz (1958) as having smaller
basidiospores (6–7 × 5.5−6.5 μm and 6–7 × 5.5−6.5 μm, respectively) and pileipellis hyphae
with clamp connections (Smith & Stuntz, 1958). Finally, P. espeletiae (Singer, 1962) is
probably the species closest morphologically to P. cf. fernandezianus due to the presence of
large basidiospores (8.0–8.8 × 6.8–7.5 µm) and cystidia (40–90 × 10–24 µm, although not
differentiated in pleuro- and cheilocystidia), but with a pileipellis described with hyphal
elements typical of a cutis: “epicutis of pileus consisting of parallel yp ae wit uppermost
members appressed to ascendant in bunches, even suberect in the region of the central
scaliness”.
Pluteus maculosipes Singer, Sydowia 15: 122, 1962.
Figs. 2d–f, 7
Pileus 25 mm diam, conic-campanulate at first and then expanding, surface greyishbrown, fibrillose-pruinose to almost pulverulent, fully covered by minute squamules, margin
not sulcate or striate. Lamellae free, at first white and then finally cream-pinkish, crowded,
ventricose, with concolourous edges and lamellulae of different lengths, 1–3 for each
lamellae. Stipe 40 × 3(apex)–5(base) mm, tapering towards the apex, central, cylindrical,
surface white to pale cream and minutely dotted brownish towards the base. Odour, taste and
flesh colour not recorded.
Basidiospores [20/1/1] 6.2–7.5 × (5.0–)5.6– .2 μm [Q = 1.10–1.24(–1.40); Qm =
1.17; Lm =
.9 μm; Wm = 5.9 μm], subglobose to broadly ellipsoid, rarely ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. Basidia 22.5–27.5 × 8.7–10.0 μm,
clavate to versiform, thin-walled, four-spored. Pleurocystidia absent. Cheilocystidia 39–56
(–74) × (6.2–)8.7–11.2(–24) μm, fusiform to ampullaceous or mucronate, usually elongated to
slender, apex rounded and usually with a narrow and short papilla, rarely longer (71–74 ×
18.7–24 μm) and broadly lageniform or slightly constricted at the mid point, colourless and
hyaline, thin-walled, very abundant in fascicules. Lamellar edge sterile. Lamellar trama
inverse, up to 38 μm wide, composed of thin-walled hyphae up to 6.2 μm diam, yaline.
Pileus context undifferentiated, appro . 50 μm t ick, composed of thin-walled hyphae up to
6.2 μm diam, yaline. Pileipellis irregular, almost a hymeniderm or a palisadoderm, with tuffs
of ascendant to erect elements from a subpellis of more or less parallel hyphae, individual
terminal elements 20–69 × 5–8.7 μm, with broadly rounded apex or sometimes tapering
Capítulo IV - 161
towards the apex, with evenly dissolved brown intracellular pigment, mainly the elements of
the subpellis, or sometimes colourless and hyaline mainly the suberect terminal elements.
Stipitipellis a cutis of thin-walled hyphae, up to 6.2 µm diam, but with many erect to suberect
cystidioid elements organized in fascicules, individual terminal elements (18.7–)29–56 × 3.7–
5.0 μm, gradually attenuated forming an obtuse to subacute apex (up to 2.5 µm diam), filled
with evenly dissolved pale straw pigment. Clamp connections absent in all parts examined.
Fig. 7. Pluteus maculosipes (FK1651). a. basidiospores; b. cheilocystidia; c. pileipellis elements;
d. caulocystidia. Scale bars: 10 µm.
Capítulo IV - 162
Habit and habitat. – Solitary, on decayed wood.
Revised specimen. – BRAZIL: Paraná State, Foz do Iguaçu city, Parque Nacional do
Iguaçu, Trilha do Poço Preto, 6 November 2010, F. Karstedt FK1651 (SP417463).
Remarks. – Pluteus maculosipes is a species described from Venezuela (Singer,
1962) and characterized by the absence of pleurocystidia, a distinct pileipellis and a
stipitipellis with accumulations of cystidioid elements organized in fascicules. Singer (1962)
reported and illustrated t e presence in lamellar sides of “pseudoparap ysis-like bodies which
are vesiculose-clavate, (… 1 .5 × 9.3 μm”, but t ese structures most likely represent
basidioles in the hymenium. In our material we observed the presence of some cheilocystidia
slightly bigger and different in shape from those commonly observed, but they are rare and
hence of doubtful taxonomic value to distinguish it from the type of P. maculosipes.
Among the few species of Pluteus with very rare to absent pleurocystidia,
P. maculosipes is different from almost all because they have a pileipellis characteristic of the
species currently classified in sect. Celluloderma, as observed in P. diettrichii, P. insidiosus
Vellinga & Schreurs, P. poliocnemis Kühner, P. rimulosus Kühner & Romagn., P. seticeps
and P. thomsonii (Berk. & Broome) Dennis. Pluteus exiguus and P. pusillulus Romagn. are
the few species lacking pleurocystidia and with a hymenidermal pileipellis of narrowly
fusiform elements or as a palisade or a trichoderm (Vellinga and Schreurs 1985, Orton 1986).
However, P. pusillulus has subcapitate cheilocystidia, pileipellis organized in a true
hymeniderm and stipitipellis without cystidioid elements (Vellinga & Schreurs, 1985), while
P. exiguus has as a distinctly squamulose pileus and clavate c eilocystidia wit a distinct
narrow ape or apical pro ections (K
ner and omagnesi 1956, Orton 1986).
Pluteus velutinus Pradeep, Justo & Vrinda, Mycol. Progress 11: 871, 2012.
Figs. 2g–i, 8–9
Pileus 35 mm diam, convex, slightly depressed at centre, surface uniformly beige to
light brown (N50A50M40), velvety to pruinose-fibrillose all over, sometimes detaching and
exposing the white translucent flesh, margin pellucid striate. Lamellae free, pinkish
(N20A20M20), moderately crowded, ventricose, up to 8 mm, with concolourous edges and
lamellulae of different lengths, 1–3 for each lamellae. Stipe 52 × 3–4 mm, slightly tapering
towards the apex, central, cylindrical, surface white-cream with brownish points on base,
fibrous, hollow, with strigose mycelium at extreme base. Odour and taste not recorded.
Basidiospores [20/1/1] (6.8–)7.5–8.7 × 5.6– .8(−7.5 μm (Q = 1.1 –1.45; Qm =
1.2 ; Lm = 7.9 μm; Wm = .2 μm , broadly ellipsoid to ellipsoid, inamyloid, yaline, smoot ,
thick-walled, guttulate. Basidia 20–26 × 6.2–7.5 μm, clavate or versiform, thin-walled, fourspored, with small scattered guttules. Pleurocystidia 62–81(−92 × (12.5–)15.0–18.7(−31
Capítulo IV - 163
μm, ventricose to utriform or lanceolate, usually wit one or two apical or lateral pro ections
like a knob-shaped appendix, rarely with a median constriction, mostly with pale straw
content, pigmentation usually concentrated at apex or sometimes evenly dissolved, rarely
colourless, thin-walled, scattered and abundant. Cheilocystidia (38–)50–61 × 10.0–15.0 μm,
ventricose to utriform or rarely clavate, with a rounded apex, colourless and hyaline, thinwalled, abundant. Lamellar edge sterile. Lamellar trama inverse, up to 31 μm wide,
composed of thin-walled hyphae, 2.5–1 .2 μm diam, interwovened by oleiferous and inflated
hyphae, hyaline. Pileus context undifferentiated, composed of thin-walled hyphae, 2.5–16.2
μm diam, interwovened by oleiferous and inflated
yp ae,
yaline. Pileipellis a
tric o ymeniderm up to 75 μm t ick, composed of clavate, vesiculose or elongate-clavate
cells, (28–)41–61 × (22–)29–34 μm, t in-walled, with brown dissolved content.
Caulocystidia 31–51 × 13.7–1 .2 μm, mostly clavate or sometimes wit
a median
constriction, thin-walled, with pale straw to brownish content. Clamp connections absent in
all parts examined.
Habit and habitat. – Subgregarious (two basidiomata near), apparently on dead leaf
sheath of palm tree.
Revised specimen. – BRAZIL: Pará State, Belém city, Ilha do Combu, 20 February
2011, F. Karstedt & C. Castro FK1889 (SP417670).
Remarks. – Pluteus velutinus was recently described from India with additional
collections from Japan (Pradeep et al., 2012) and appears in the phylogenetic analyses in the
plautus clade as sister to P. longistriatus (Fig. 1). The ITS sequence of P. velutinus from
Brazil is up to 99.47% identical (three base pair differences) with the sequence of the
holotype and up to 99.65% identical (two base pair differences) with the Japanese sequences.
In addition, a new sequence of P. velutinus from Mongolia is also included in the analyses
(Fig. 1), confirming the broad geographic distribution for the species.
Comparing the Brazilian collection to the Asian materials, some minor
morphological differences were observed, mainly regarding the shape and size of the
microstructures and the most evident difference is the pigmentation and the apical shape of
pleurocystidia. The majority pleurocystidia of the Brazilian collection has a pale straw content
and apical or lateral projections like a knob-shape, while those studied by Pradeep et al.
(2012) are colourless and commonly with an apical digitate projection at the apex.
Also, the cheilocystidia herein studied are more slender than those described in the
protologue (up to 30 μm widt
and preponderantly wit a rounded ape , like t ose illustrated
from the tropical collection (holotype from India) but different from those with a small apical
projection that are also characteristic for P. velutinus, but apparently restrict to the Japanese
Capítulo IV - 164
collections as illustrated by Pradeep et al. (2012). Finally, the pileipellis cells of the Brazilian
collection are much smaller in length and the caulocystidia are slightly more slender than
those described on the protologue that are up to 140 μm in lengt for t e pileipellis cells and
up to 30 μm in widt for t e caulocystidia.
Fig. 8. Pluteus velutinus (FK1889). a. basidiospores; b. pleurocystidia. Scale bar: 10 µm.
The occurrence of P. velutinus in Brazil and India shows an apparent pantropical
distribution for this species. However, its occurrence in a temperate area in Japan (Hokkaido)
is an exception for this pattern. Data from the nearest climatological station (Iwamizawa
Station – Japan Meteorological Agency) indicated that during the period of fructification (July
and September 2005) of the collections studied by Pradeep et al. (2012), the monthly mean
Capítulo IV - 165
temperature was respectively 19.2°C (daily mean min. 15.7°C and max. 24°C) and 17.5 °C
(daily min. 12.5°C and max. 22.7°C), while the monthly mean relative humidity was 79 and
77%, respectively. These data show a considerable difference between the climatic conditions
observed during the preceding period of collecting in Brazil. Data from a climatological
station installed in the collecting area (Ilha do Combu) indicated that during the beginning of
the rainy season (mid-December 2010 and mid-January 2011) the mean temperature was
24.9°C (daily min. 22.0°C and max. 30.9°C) while the mean relative humidity was 90.62%
(Freire, 2011 and pers. com.).
Fig. 9. Pluteus velutinus (FK1889). a. cheilocystidia; b. pileipellis elements; c. caulocystidia. Scale bars: 10 µm.
Capítulo IV - 166
As previously pointed out by Pradeep et al. (2012), it is difficult to understand the
distribution pattern of P. velutinus in both temperate and tropical zones with different climatic
conditions and habitat characteristics and whether its dispersal is natural or anthropic. On the
other hand, the coexistence of P. velutinus in similar but distant tropical areas such as Brazil
and India could be better understood if we consider the general climatic conditions between
both localities. However, the collecting area of P. velutinus in Brazil represent a very
interesting habitat that, although it being part of a tropical region, is located in a particular
area of the Amazonian forest that is extremely peculiar to fungal adaptation and fructification
due to the condition of periodical inundations (Singer, 1984).
This type of Amazonian forest is known as várzea (swamp) and it is characterized by
a seasonal variation of flooding caused by white-water rivers (Prance, 1979). The area
occupied by the Ilha do Combu can be considered a specific type of várzea called Tidal
várzea forest, which is flooded and drained twice daily by the tidal movements, because high
tides temporarily block the flow of rivers in the estuarine region and cause them to flood
towards the adjacent forest (Prance, 1979).
The periodical várzea inundation causes deficiency of the O2 level that affects
directly the fungal development and also prevents the basidiospores discharge at ground level
(Singer, 1984). In addition, according to Singer (1984), the várzea forest is clearly
anectotrophic (free of ectomycorrhiza) what precludes an alternative mycelial survival
strategy restricted to ectomycorrhizal and root parasitic fungi. So that, Singer (1984) proposed
four different ways to understand how the saprophytic and parasitic basidiomycetes adapt
themselves to a long and deep immersion under várzea conditions.
According to the period and pattern of fructification observed for P. velutinus from
Brazil and considering the proposition of Singer (1984), probably the adaptation of
P. velutinus is related to “some of the litter and alluvium inhabiting, mostly smaller
Agaricales form large numbers of carpophores in the relatively short period between the
beginning of the rainy season in December and the time when inundation cover the ground
level. It is remarkable that many of these produce ± thick-walled spores or mycelia capable of
remaining viable during several months of immersion with the current lack of oxygen”.
Considering that P. velutinus occurs in so distant tropical areas (Brazil and India) and also
that it is able to develop something like this survival strategy, maybe a similar adaptation
pattern could be used to explain its occurrence in temperate conditions and lower
temperatures.
Within this same adaptation pattern, Singer (1984) reported a species apparently
endemic to várzea, which was later described (Singer, 1989) as P. varzeicola from Amazonas
Capítulo IV - 167
State, Brazil. Although P. velutinus and P. varzeicola apparently share the same adaptation
pattern for resistance to várzea conditions, the re-examination of the holotype of P. varzeicola
(BRAZIL: Amazonas State, Ilha Marchantaria, 9 March 1983, Singer B 12397, F!) indicated
that they are very close in the morphological concept.
The holotype of P. varzeicola (Fig. 10, 11) has broadly ellipsoid to ellipsoid or rarely
subglobose basidiospores [20/1/1] 6.2–7.5(–8.1) × 5.0–6.2(–6.8) μm (Q = 1.10–1.50; Qm =
1.30; Lm = 7.2 μm; Wm = 5.5 μm ; clavate to ventricose or lanceolate pleurocystidia, 45–62
× 13.7–21 μm, w ic are wit out any kind of pro ections and mostly wit concentrated pale
straw content or rarely colourless; clavate to vesiculose and colourless cheilocystidia, 32–57 ×
15.0–27 μm; tric o ymenidermic pileipellis composed of vesiculose, clavate to fusoid cells
with dissolved or commonly condensed brown content, (43–)60–90(–112) × 18.7–34 μm; and
clavate to fusoid caulocystidia, 44–87 × 13.7–20 μm, w ic are colourless and rarely with
narrow papillate apex.
Fig. 10. Pluteus varzeicola (Singer B 12397 – holotype). a. basidiospores; b. pleurocystidia; c. cheilocystidia.
Scale bar: 10 µm.
It is a hard matter to distinguish P. velutinus from P. varzeicola since the
micromorphology of both species is basically the same, except because P. varzeicola has
Capítulo IV - 168
pleurocystidia that are completely devoid of any apical projections, colourless caulocystidia
and more broadly clavate and slightly smaller cheilocystidia, 15–50 × 10–24 µm measured by
Singer (1989) and 32–57 × 15.0–27 µm measured after our holotype re-examination. In the
macromorphology, P. varzeicola differs from P. velutinus in having a pileus with the slightly
sulcate margin and slightly venose-rugose at centre (Singer 1989).
Fig. 11. Pluteus varzeicola (Singer B 12397 – holotype). a. pileipellis elements; b. caulocystidia. Scale bars: 10
µm.
Regarding the pleurocystidia pigmentation, the collection of P. velutinus from Brazil
has both pigmented and colourless pleurocystidia as also P. varzeicola, but P. velutinus from
Capítulo IV - 169
India and Japan has only colourless pleurocystidia. The caulocystidia pigmentation is a
common character that is shared between all collections of P. velutinus (from Asia and South
America), but absent in P. varzeicola. Finally, the measurements for P. velutinus
cheilocystidia are 38–61 × 10.0–15.0 µm in this paper (from FK1889) and 32–70 × (9.5–)
12–30 µm by Pradeep et al. (2012).
Despite these micromorphological differences, which in part can be also observed
between Indian and Japanese collections according to illustration from Pradeep et al. (2012),
and also when they are compared to the Brazilian collection, just based on molecular results
(Fig. 1) it was possible to recognize them as the same species with a relatively wide range of
morphological variation and mainly a considerable habitat divergence over the widespread
geographic distribution.
Considering that all collections involving P. varzeicola and P. velutinus have
individual micromorphological differences, it is difficult to establish any distinguishable
feature to separate or to recognize reliably them as the same taxa, except for the pileus
characters. Thus, although we suspect that P. velutinus could be considered synonym of
P. varzeicola, until more molecular data from Pluteus inhabiting the várzea habitat could be
obtained, we prefer consider them as very closer in the morphological concept and maintain
this record under P. velutinus expanding its geographical distribution.
Acknowledgements. The authors thank the curators of F, RSPF and SP for the loan of the specimens
studied; Antonio Sérgio Cunha Freire for providing the meteorological data from Ilha do Combu;
Noemia Kazue Ishikawa and Hitoshi Neda for helping with the meteorological data from Hokkaido;
Fernanda Karstedt for collecting the specimen and providing the photographs used in this study; Helen
Maria Pontes Sotão for helping during the expedition of F. Karstedt to Ilha do Combu in Pará State;
Flávia Ribeiro Santos for helping with DNA extraction of some samples; Klei R. Sousa for inking the
line drawings. N. Menolli r. and M. Capelari t ank “Funda o de Amparo à Pes uisa do Estado de
S o Paulo” (FAPESP grants 04/04319-2, 09/53272-2 , “Coordenação de Aperfeiçoamento de Pessoal
de N vel Superior” (CAPES - PNADB grant 23038.000043/2010-43 and t e “Consel o Nacional de
Desenvolvimento Cient fico e Tecnológico” (CNP
fellows ips for financial support. A. Justo
acknowledges support from the National Science Foundation grant DEB 0933081.
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CAPÍTULO V
Pluteus sect. Celluloderma in Brazil:
phylogeny and taxonomy including eight new species *
Nelson Menolli Jr., Alfredo Justo & Marina Capelari
*Artigo a ser submetido para publicação na revista Mycologia
Capítulo V - 173
Capítulo V - 174
Short title: Pluteus sect. Celluloderma in Brazil
Pluteus sect. Celluloderma in Brazil: phylogeny and taxonomy including eight new species
Nelson Menolli Junior
CCT/Biologia, Rua Pedro Vicente 625, 01109-010, São Paulo, SP, Brazil, and Núcleo de
Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902, São Paulo,
SP, Brazil
Marina Capelari
Núcleo de Pesquisa em Micologia, Instituto de Botânica, Caixa Postal 68041, 04045-902,
Alfredo Justo
Clark University, Biology Department, 950 Main St., Worcester, 01610 MA, USA
Abstract: A general phylogeny of Pluteus sect. Celluloderma based on ITS sequences is
presented with description of eight new species based on morphological and molecular data:
P. aureolatus, P. brunneocrinitus, P. cebolinhae, P. crinitus, P. hispidulussimilis,
P. karstedtianus, P. necopinatus and P. paucicystidiatus. A revision of Pluteus sect.
Celluloderma occurring in Brazil is also presented confirming the occurrence of 29 taxa
considered certainly known for the country.
Key words: Agaricales, biodiversity, ITS, Pluteaceae
Capítulo V - 175
INTRODUCTION
The traditional subdivision of Pluteus into three sections (Pluteus, Celluloderma Fayod and
Hispidoderma Fayod) has been supported by recent molecular phylogenetic studies (Menolli
et al. 2010, Justo et al. 2011a, b) with some rearrangements. Sect. Celluloderma, as
recognized by Singer (1959, 1986), includes species with non-metuloid pleurocystidia and
pileipellis composed of a hymeniform layer of relatively short elements, interrupted or not by
elongate cystidioid bodies. Singer (1956, 1959, 1986) further subdivided sect. Celluloderma
into two subsections: Mixtini Singer, characterized by the presence of elongated elements in
the pileipellis, and Eucellulodermini Singer, characterized by the lack of these elements.
Vellinga and Schreurs (1985) proposed a distinct organization for this classification
system subdividing the sect. Celluloderma into three subsections: Mixtini, Eucelullodermini
and Hispidodermini (Fayod) Vellinga & Schreurs, which is characterized by a trichodermal
pileipellis or by a hymeniderm of cylindrical to fusiform elements that corresponds to part of
the members of sect. Hispidoderma sensu Singer (1959, 1986). The species characterized as
having a cutis-like pileipellis were accommodate in a new section, viz. sect. Villosi Schreurs
& Vellinga, which was introduced for the remaining members of sect. Hispidoderma sensu
Singer (1959, 1986). However, recent molecular phylogenetic studies including species of
sect. Celluloderma (Justo et al. 2011a, 2012) do not support the division in subsections
proposed by Singer (1956, 1959, 1986) or Vellinga and Schreurs (1985).
Although these different rearrangements, based on molecular data (Minnis et al.
2006, Menolli et al. 2010, Justo et al. 2011a, b, Vizzini and Ercole 2011) sect. Celluloderma
now includes species with non-metuloid pleurocystidia and pileipellis composed of a
hymeniform layer or as an epithelium of relatively short elements, interrupted or not by
elongate cystidioid bodies (corresponding to sect. Celluloderma as defined by Singer 1956,
1959, 1986) and also species with a cutis-like pileipellis (corresponding to sect. Villosi or
Hispidoderma pro parte) or annulate species (corresponding to the genus Chamaeota).
After a morphological and molecular study of new collections of Pluteus sect.
Celluloderma, we provide a phylogenetic overview for this section with special emphasis to
species occurring in Brazil, including new records and new species, and presenting a list of all
species previously recorded for the country with an update of the taxonomic status of the
Capítulo V - 176
MATERIALS AND METHODS
Sampling.
The materials studied in the morphological and molecular analyses include specimens recently collected in
Brazil and collections previously published and/or deposited in herbaria. Searches of the bibliographical and
herbarium records were the basis to prepare a list with the current knowledge of sect. Celluloderma in Brazil.
T e erbarium acronyms follow T iers (2013 and t e ‘ ede
rasileira de Herb rios’ ( ttp://www.botanica.
org.br/rede_herbarios.php) for the Herbarium from Passo Fundo University (RSPF).
Species characterized as having non-metuloid pleurocystidia and a pileipellis as a cutis were included
in this work because we accept the section delimitation as complemented by Justo et al. (2011a, b) that
considered these species in sect. Celluloderma instead of in sect. Hispidoderma or Villosi as recognized by
Singer (1959, 1986) and Vellinga and Schreurs (1985), respectively.
Molecular study and phylogenetic analyses.
Methods for DNA isolation, PCR and sequencing follow Justo et al. (2011b). The ITS region was amplified
using the primer pair ITS1-F and ITS4 (White et al. 1990, Gardes and Bruns, 1993). The molecular analyses
were conducted with existing DNA sequences used by Menolli et al. (2010), Justo et al. (2011a, b, c, 2012),
Pradeep et al. (2012) and from newly collected sequences of materials representing species of sect.
Celluloderma. Newly obtained sequences were deposited in GenBank. All GenBank accession or collection
numbers and sequence geographic origins are given in Fig. 1. Sequences were aligned using MAFFT version 6
(http://mafft.cbrc.jp/alignment/server/; Katoh and Toh 2008) with the Q-INS-i option. Then, the alignment was
visually examined and manually corrected using MacClade 4.05 (Maddison and Maddison 2002). Maximum
Parsimony (MP) and Maximum Likelihood (ML) were performed with the parameters specified in Justo et al.
(2011b). Pluteus cervinus (Schaeff.) P. Kumm. and P. petasatus (Fr.) Gillet were used as outgroup taxa.
Sequence divergence was calculated using MatGAT (Campanella et al. 2003).
Morphological study.
The macroscopic description was based on fresh specimens. Color terms are according to Küppers (1979) for all
descriptions except for P. paucicystidiatus that follows Kornerup and Wanscher (1978). In some cases the
macroscopic description of the Brazilian collections was not recovered but other references are provided below
the taxonomic entry to complement the species recognition. For microscopic analyses, the dried material was
wetted with 70% ethanol and then rehydrated in 5% KOH or stained wit Melzer’s reagent to determine t e
amiloidity reaction of the basidiospores. The notation [a/b/c] at the beginning of a set of basidiospore data is to
be read as “(a basidiospores were measured from (b basidiomata taken from (c collections”. Q represents the
range of the length/width ratio for all of the measured spores, Qm represents the average of all calculated Q
values for all of the measured basidiospores and Lm (Wm) represents the average of all of the lengths (widths) of
the measured basidiospores. At least 20 basidiospores from each basidioma were measured in lateral view, and
the terms denoting basidiospore shape follow Bas (1969).
Capítulo V - 177
RESULTS
The final dataset consists of 154 ingroup sequences with a total of 799 characters (gaps
included), of which 394 are parsimony informative. In the MP analysis 146 MPT were
recovered (Length = 2698; CI = 0.31; RI = 0.84). A general phylogeny of sect. Celluloderma
based on the best tree from the ML analysis is shown in Fig. 1.
In both ML and MP analyses seven well-supported clades were recognized:
aurantiorugosus clade ( S ≥ 70 only in MP analysis , cinereofuscus clade, ephebeus clade,
podospileus clade, hispidulus clade, thomsonii clade and phlebophorus clade. Three minor
clades formed by pairs of sequences were also recognized: i) P. brunneosquamulosus C.K.
Pradeep & K.B. Vrinda and Pluteus eliae Singer; ii) P. hispidulus (Fr.) Gillet
(ARAN8200509) and Pluteus sp. (AJ588); iii) P. cebolinhae and P. crenulatus Justo, Battistin
& Angelini. Additionally, P. diettrichii Bres. appears external to all taxa in sect. Celluloderma
except of P. cebolinhae and P. crenulatus. Although the recognition of these clades, the
relationship between then are not resolved because different rearrangements were verified in
previous studies including molecular analyses of sect. Celluloderma species (Justo et al.
2011a, b, c, 2012, Pradeep et al. 2012).
As previously pointed out by Justo et al. (2011a) and confirming the artificial status
of the subsections proposed by Singer (1956, 1959, 1986), taxa with Mixtini-type pileipellis
are gathered in the podospileus and thomsonii clades, which are primarily formed by species
with this type of pileipellis (except by P. dominicanus var. hyalinus Menolli & Capelari and
other sequences from which the morphological data are unavailable until now, viz. JX545226,
AJ838 and AJ491), but also spreaded in at least three other clades: aurantiorugosus clade
(P. stenotrichus Justo, Battistin & Angelini), cinereofuscus clade (P. eludens E.F. Malysheva,
Minnis & Justo and P. multiformis Justo, A. Caball. & G. Muñoz) and phlebophorus clade
(P. cf. eugraptus). The occurrence of a cutis-like pileipellis also evolved many times during
the evolution of sect. Celluloderma because taxa with this character appeared at least in three
independent clades: aurantiorugosus clade [P. cubensis (Murril) Dennis], ephebeus clade (all
taxa) and hispidulus clade (all taxa).
The species identification of many non-Brazilian sequences presented in the tree
must be revised and some other yet to be identified, but it falls outside the scope of this paper.
All of these sequences are marked with an asterisk and previous indications of some probable
identification can be seen in Justo et al. (2012).
Based on morphological and molecular data we propose eight new species:
P. aureolatus, P. brunneocrinitus, P. cebolinhae, P. crinitus, P. hispidulussimilis,
P. karstedtianus, P. necopinatus and P. paucicystidiatus. The taxonomy and phylogenetic
Capítulo V - 178
position of P. diptychocystis, P. iguazuensis and P. rimosoaffinis are also discussed. The
record of P. cf. fastigiatus and P. cf. fuliginosus is maintained as uncertain determination due
to some ecological and micromorphological differences between the type collections (Murrill
1917, Singer 1959) and our materials.
Fig. 1. Best tree from the maximum likelihood analysis for the ITS dataset of Pluteus sect. Celluloderma.
ootstrap values ≥70 % are s own on branc es (ML/MP . oot lengt
as been reduced to facilitate grap ical
representation. *** Voucher collections that need revision for correct identification. # # # Voucher collections
that were re-identified (see discussion on text).
Capítulo V - 179
Fig. 1. Concluded.
The current knowledge of sect. Celluloderma in Brazil, as accepted in this paper, is
presented in Table 1. Searches of the bibliographical and herbarium records, complemented
by the species herein presented, led to a total of 54 taxa names of sect. Celluloderma that
were linked to specimens collected in Brazil (Table 1). However, only 29 taxa are here
considered certainly known from Brazil, with 21 of them phylogenetically positioned in the
current analyses. Additionally, the records of P. amazonicus Singer, P. fallax Singer and
P. paraensis Singer need new collections to certify their existence in Brazil (Table 1). The
occurrence in Brazil of the other 22 taxa were not confirmed because the collections
associated to these names were not located, are too insufficient for study, or represent species
of other sections. Data of previous studies were also used to endorse the occurrence of these
taxa in Brazil (Singer 1959, 1973a, 1989, Menolli and Capelari 2010, 2013, Menolli et al.
2010, 2013b).
Capítulo V - 180
The infrageneric position of some species in sect. Celluloderma should be regarded
with caution. The position of P. amazonicus is discussed by Singer (1959) as doubtful
between sections Hispidoderma and Celluloderma due to its pileipellis structure with
elements varying from short to long. However, considering the average of length/width ratio
(Qm < 3, according to Justo et al. 2011a) for the measuments of the pileipellis elements done
by Singer (1989), we consider P. amazonicus better classified in sect. Celluloderma.
Although the Brazilian type was not located at INPA, we prefer to maintain this species as
occurring in Brazil based on the complete description presented by Singer (1989).
It is not clear to us the real position of P. minutus, but based on the characteristics of
the pileipellis described by Singer (1956) we prefer to maintain it in sect. Celluloderma. It is
important to note that although Singer (1989) mentioned the collection from Bahia State,
Itabuna (R. Singer B 11119, 7 jun. 1973, INPA) as P. minutus, but it is deposited at INPA as
P. oligocystis var. dennisii (Singer) Singer and dated of 6 Jun. 1978, which probably is the
correct date according to the field itinerary of R. Singer (Strack and Mueller 1997).
Nevertheless, its occurrence in Brazil could not be confirmed because the material available
in INPA is too insufficient to study and because R. Singer did not provide any description for
this collection.
Pluteus avellaneus, P. hispidulus and P. pluvialis are considered species having a
cutis-like pileipellis and so listed as species of sect. Celluloderma previously recorded from
Brazil, but only the occurrence of P. pluvialis is confirmed (Table 1).
Table 1: Species of Pluteus sect. Celluloderma recorded from Brazil and notes about the identification of the
Species
Reference or herbarium data
P. amazonicus Singer
Singer (1989)
P. anomocystidiatus Menolli & de Menolli et al. (2013b)
Meijer
P. atriavellaneus Murrill
unpublished data (SP233830)
P. aureolatus
P. aureovenatus Menolli &
Capelari
P. avellaneus Murrill
P. beniensis Singer
Notes about identification
of the Brazilian collections
unavailable for study
P. glaucotinctus E. Horak
(Menolli et al. 2013a)
this paper
P. brunneocrinitus
P. brunneopictus Berk. & Broome
unpublished data (SMDB9193)
Stijve & Meijer (1993 as P. cf. beniensis)
Meijer (2006 as P. cf. beniensis)
this paper
Rick (1938, 1961*)
P. burserae Singer
P. cebolinhae
Singer (1959)
this paper
not found
most likely P. tucumanus
Singer and P. cubensis
(Menolli and Capelari
2013)
Capítulo V - 181
Table 1. Continued.
Species
P. chrysophlebius subsp. bruchii
(Speg.) Singer
P. crassus Menolli & de Meijer
P. crinitus
P. cubensis
Meijer (2006)
this paper
Stijve & Meijer (1993)
Pegler (1997*)
Meijer (2006*)
Rosa et al. (2006)
Xavier-Santos et al. (2004 as P. cf.
cubensis)
Rosa & Capelari (2009)
P. diptychocystis
P. dominicanus var. hyalinus
P. eludens
P. eugraptus (Berk. & Broome)
Sacc.
Justo et al. (2011a, b as Pluteus sp. V)
Pegler (1997)
this paper
Meijer (2006)
P. fallax Singer
P. cf. fastigiatus Singer
Singer (1959)
this paper
P. fluminensis
Singer (1959)
Stijve & Meijer (1993)
Meijer (2006 as P. cf. fluminensis)
Pegler (1997)
P. fuligineovenosus E. Horak
P. cf. fuliginosus Murrill
this paper
P. fusconigricans (Berk. &
Broome) Sacc.
P. globiger Singer
P. hispidulus
P. hispidulussimilis
P. homolae Minnis & Sundb.
P. iguazuensis
this paper
Meijer (2006 as P. cf. iguazuensis)
P. jamaicensis
Drechsler-Santos et al. (2007)
this paper
P. karstedtianus
P. laetifrons var. laetifrons (Berk.
& M.A. Curtis) Sacc.
P. laetifrons var. floridae Singer
Rick (1919, 1938, 1961)
this paper
Meijer (2006, 2008)
Meijer (2006)
P. crassus (Menolli et al.
2013b)
P. cubensis and P.
albostipitatus (Menolli et
al. 2013a, b)
P. cubensis, P. rimosellus
and P. striatocystis
(Menolli et al. 2013b)
P. argentinensis (Menolli et
al. 2013b)
P. argentinensis (Menolli et
al. 2013b)
P. rimosoaffinis (this paper)
P. chusqueae (E. Horak)
Menolli (Menolli et al.
2013b)
not found
see comments in Taxonomy
part
P. riograndensis (Menolli
et al. 2013b)
et al. 2013b)
et al. 2013b)
et al. 2013b)
see comments in Taxonomy
part
most likely P. yungensis
Singer (Menolli and
Capelari 2013)
P. crassus (Menolli et al.
2013b)
et al. 2013b)
insufficient for
identification
not found
Capítulo V - 182
Table 1. Concluded
Species
Rick (1919, 1938, 1961)
P. nanus (Pers.) P. Kumm.
P. nanus var. podospileus (Sacc. & Rick (1938, 1961)
Cub.) Rick
P. aff. neophlebophorus
Meijer (2006)
P. microsporus
unpublished data (SP250534)
P. minutus Pat.
P. necopinatus
P. paraensis Singer
P. paucicystidiatus
Singer (1989)
this paper
Singer (1973a)
Justo et al. (2011a, b as Pluteus sp. IV)
this paper
Rick (1938, 1961*)
P. phlebophorus (Ditmar) P.
Kumm.
P. pluvialis Singer
Pegler (1997)
P. pulverulentus var. pseudonanus Meijer (2006)
Singer
P. riberaltensis var. conquistensis Menolli et al. (2010)
Singer
P. rimosoaffinis
Pegler (1997)
Meijer (2006* as P. cf. rimossoaffinis)
P. riograndensis Singer
P. sapiicola Singer
P. striatocystis Pegler
P. sublaevigatus (Singer) Menolli
& Capelari
P. thomsonii (Berk. & Broome)
Dennis
Justo et al. (2011a, b as Pluteus sp. III)
this paper
Singer (1954b, 1956, 1959)
Raithelhuber (1991)
Justo et al. (2011a,b as Pluteus sp. II)
Singer (1954b, 1956, 1959)
Raithelhuber (1991)
Meijer (2006, 2008)
Rosa & Capelari (2009)
Menolli & Capelari (2010)
Singer (1956)
P. variipes Singer
Names in bold are considered species certainly known in Brazil.
classification of these species.

P. eludens (Menolli et al.
2013b)
not found in herbarium
collections
not found
most likely P. sapiicola and
P. tucumanus (Menolli
and Capelari 2013)
P. longistriatus (Menolli et
al., 2013a)
not found
P. riograndensis and P.
sapiicola (Menolli et al.
2013b)
Meijer (2006 as P. cf. thomsonii)
P. umbrinoalbidus
most likely P. sapiicola
(Menolli and Capelari
2013)
not found

P. thomsonii (Menolli et al.
2013b)
not found
re-identified as P.
fluminensis by Singer
(1959)
= See discussion above about the infrageneric
= Species that need new collections to certify their existence in Brazil.

= Species not
studied in this work but that are considered as occurring in Brazil due to the well-preserved collections and the complete
description presented by the authors referenced.

= Reference without indication of the collection studied and/or the
herbarium where the materials were deposited. * = Reference that includes two or more collections that are actually different
species.
Capítulo V - 183
Although the occurrence of P. variipes in Brazil was disregarded because Singer
(1959) re-identified the Brazilian collection (R. Singer B 432) as P. fluminensis, we located in
FH a collection of P. variipes (R. Singer B 1425 – FH00301675) presumably from Brazil
according to the exsiccatum label. However, this collection obviously is not from Brazil but
actually from Bolivia because Singer (1959) mentioned R. Singer B 1425 as a specimen of
P. variipes collected in Bolivia on 23 Feb. 1956. Moreover, according to Strack and Mueller
(1997), during 1956, R. Singer arrived in Brazil only in March. Collections of other species
made on 23 Feb. 1956 (Pluteus oligocystis Singer, R. Singer B 1420 and Crepidotus
yungicola Singer, R. Singer B 1422) were also referenced from Bolivia (Singer 1959, 1973b,
Wu et al. 1997), confirming the mistake available in FH. Thus, the occurrence of P. variipes
from Brazil can be disregarded.
Additional molecular data are still needed to certify the recognition of some
Brazilian collections in the aurantiorugosus and cinereofuscus clades. The sequences within
the P. aureovenatus complex (Fig. 1) present values of sequence divergence varying between
0.20% and 4.58%. Current studies using combined analyses of ITS + TEF1 (data not show)
revealed the existence of at least four putative species in this complex. Thus, for now we
confirm the occurrence of P. aureovenatus (FK1699 and FJ816663), P. globiger (JQ065030)
and P. sublaevigatus (FJ816667) from Brazil, but keeping as Pluteus sp. the collection
FK1602 and safeguarding the identification of the sequence HM562160.
A further multi-locus study also could indicate the existence of different species
within the same broad morphological concept of the collections within the P. riograndensis
complex (Fig. 1). All of these sequences present values of sequence divergence varying
between 0.17% and 3.26%, with the higher values occurring between FJ816664/FJ816665
and all others. Despite this molecular divergence, all collections share the same
morphological concept applied to P. riograndensis. Some collections in this complex were
previously identified (Menolli et al. 2010) as P. fluminensis (FJ816664 and FJ816665),
P. fuligineovenosus (FJ816662) and P. jamaicensis (FJ816657). However, a recent
morphological re-examination (Menolli et al. 2013b) of these specimens showed that they
better fit the concept of P. riograndensis by the presence of pigmented cheilocystidia,
although they are intermixed by some colorless. Hitherto now we prefer to maintain all
collections in this complex as P. riograndensis sensu lato and so excluding the record of
P. jamaicensis and P. fuligineovenosus from Brazil (Table 1), but keeping the occurrence of
P. fluminensis by the type record (Singer 1959).
Capítulo V - 184
TAXONOMY
Pluteus aureolatus, Menolli, Justo & Capelari, sp. nov.
FIGS. 2a, b, 3
Etymology. The name refers to the apical cover present on cystidia that resembles the
representation of the golden halos of some saints and angels.
Pileus 29–30 mm diam, plane to finally plane-concave, with a distinct umbo, surface
brown (N80Y50M40) covered by appressed and radially arranged fibrils, sometimes exposing at
margin the pinkish and translucent background between the fibrils, margin not striate or
sulcate. Lamellae free, pinkish (N10Y50M40), close to subdistant, ventricose, approx. 4 mm
broad, with concolourous edges and few lamellulae. Stipe 40–43 × 3–4(apex)–5–6(base) mm,
attenuated upwards, base sometimes clavate to subbulbous, central, surface brownish
concolour with the pileus, covered by brown fibrils overall on pale background, with or
without scanty basal mycelium. Odor, taste and flesh color not recorded.
Basidiospores [40/2/2] 5.6–6.2 × 5.6– .2 μm (Q = 1.00–1.11; Qm = 1.03; Lm = 6.1
μm; Wm = 5.9 μm , globose to subglobose, inamyloid, hyaline, smooth, thick-walled,
guttulate. Basidia 18.7–26 × 7.5–8.7 μm, clavate to ventricose, thin-walled, four-spored.
Pleurocystidia 42–55 × 13.7−21(–32) μm, fusoid-ventricose to lageniform, colorless and
without internal content, hyaline, thin-walled, mostly with an apical yellowish resinous
incrustation forming a rigid spherical outline at the tip, sparse and moderately abundant.
Cheilocystidia (32–)42–58 × 13.7–34 μm, similar to the pleurocystidia but mostly without the
apical cover and frequently cylindrical to inflated clavate or with basal septum, colorless and
hyaline, thin-walled, numerous. Lamellar edge sterile. Lamellar trama inverse, up to 31 μm
wide, composed of thin-walled hyphae, up to 10.0 μm diam, hyaline. Pileus context
undifferentiated, approx. 150 μm t ick, composed of t in-walled hyphae, 2.5–15.0 μm diam,
hyaline. Pileipellis a cutis up to 125 μm t ick, composed of parallel chains of cylindrical and
thin-walled hyphae, individual terminal elements 112–126 × 20–2
μm, sometimes few
elongated, ascendant and conspicuous subradial elements, mostly with rounded apex or
gradually attenuated towards the apex (6.0–10.0 μm broad), sometimes with incrusted wall at
the apex, filled with evenly dissolved brownish intracellular pigment or rarely with colorless
terminal elements. Caulocystidia absent. Stipitipellis composed of thin- to thick-walled
hyphae, 6.2–18.7 µm diam, yellowish or with brownish content mainly in the terminal
elements. Clamp connections absent in all parts examined.
Habit and habitat. Solitary on soil.
Specimens examined. BRAZIL. AMAZONAS: Manaus, Instituto Nacional de
Pesquisas da Amazônia (INPA), Campus Aleixo I, 29 Jun 2011, F. Karstedt FK2084
Capítulo V - 185
(INPA241273 – HOLOTYPE); 9 Feb. 2011, N.K. Ishikawa & D. Komura NKI7
(INPA239965).
Fig. 2. a–b. Pluteus aureolatus (FK2084 – holotype). c, f. P. brunneocrinitus (MC4535 – holotype). d–e.
P. cebolinhae (CAC39 – holotype). g–h. P. crinitus (FK2064 – holotype). i. P. diptychocystis (NMJ184).
j. P. cf. fastigiatus (NKI12). k–l. P. cf. fuliginosus (FK2158). m–n. P. hispidulussimilis (NMJ179 – holotype).
o–p. P. karstedtianus (FK1700 – holotype). q–r. P. necopinatus (FK1701 – holotype). Bars: a, b, d, e, i–l, q, r =
1 cm; c, g, h, m–p = 5 mm; f = 1 mm.
Capítulo V - 186
Fig. 3. Pluteus aureolatus (FK2084 – holotype). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia.
d. Pileipellis elements. Bars = 10 µm.
Comments. Pluteus aureolatus is characterized by the brownish stipe covered by
fibrils overall, globose to subglobose basidiospores, pleurocystidia covered by an apical
yellowish resinous incrustation forming a rigid spherical outline at the tip, and the humicolous
habitat.
Among the species with a cutis-like pileipellis and cystidia with an apical cover,
none of them grows on soil, but considering the other morphological characters P. aureolatus
is close to P. circumscissus Singer, P. glyphidatus (Berk. & Broome) Sacc. and
P. riberaltensis var. missionensis Singer. However, P. circumscissus is distinguished by the
Capítulo V - 187
presence of a narrow volval belt at the stipe base, the short-ellipsoid and non-globose
basidiospores, and by the shape (mucronate and ampullaceous) and pigmentation of the
pleuro- and cheilocystidia (Singer 1959). Pluteus glyphidatus, as reported by Pegler (1986),
has a whitish to pale yellow and glabrous stipe and caespitose clusters of dermatocystidioid
elements on the cutis pileipellis. Finally, P. riberaltensis var. missionensis, which probably is
the morphologically closer species to P. aureolatus, has a pure white stipe without any dark
fibrils (Singer 1962) as the main characteristic to separate them. Singer (1959, 1962)
described two varieties for P. riberaltensis Singer, viz. P. riberaltensis var. conquistensis
Singer and P. riberaltensis var. missionensis. The first is different from the type variety by the
presence of black stripes on the stipe (Singer 1959, Menolli et al. 2010) and the second due to
the apical cover on cystidia. Although the morphological similarities between the two
varieties of P. riberaltensis with P. aureolatus, it is here proposed as a new species by
combining these both distinctive characters and the molecular results.
Although this new taxon resembles morphologically P. riberaltensis and its varieties,
the molecular analyses (Fig. 1) showed that P. aureolatus is phylogenetically more closely
related to P. fenzlii (Schulzer) Corriol & P.-A. Moreau, P. mammillatus (Longyear) Minnis,
Sundb. & Methven, P. cf. fastigiatus, P. cf. fuliginosus and other two undetermined
collections from Dominican Republic (AJ535) and the Virgin Islands (AJ478).
Pluteus brunneocrinitus, Menolli, Justo & Capelari, sp. nov.
FIGS. 2c, f, 4
Etymology. The name refers to the morphological and phylogenetic relationship with
Pluteus crinitus in addition to the distinctive characters including the brownish punctations at
stipe, the brownish lamellar edge and the pigmented cheilocystidia.
Pileus 6–12 mm diam, convex then applanate, dark brown (N80Y90M60) and even
darker at center or in mature specimens, surface fully covered by brownish erected fibrils,
densely tomentose-fibrillose to spinulose, sometimes cracking and showing the whitish flesh,
margin not striate or sulcate but with tooth-shaped scales hanging over the edge. Lamellae
free, pinkish, subdistant, ventricose, with brown lamellar edge and 1–3 lamellulae for every
one series of lamellae. Stipe 12–25 × <1(apex)–1(base) mm, slightly attenuated upwards, with
a small subbulbous base, central, surface translucent-white, covered by small brownish fibrils
overall, apparently without basal mycelium. Odor, taste and flesh color not recorded.
Basidiospores [20/1/1] 4.3–5.6 × 4.3(–5.0 μm [Q = 1.00–1.16(–1.30); Qm = 1.13;
Lm = 5.0 μm; Wm = 4.4 μm , globose to broadly ellipsoid, inamyloid, hyaline, smooth, thickwalled, guttulate. Basidia 20–26 × 6.2–8.7 μm, clavate to ventricose-clavate, thin-walled,
four-spored. Pleurocystidia absent. Cheilocystidia 41–66 × 13.7–16.2(–22) μm, elongate-
Capítulo V - 188
clavate, filled with evenly dissolved brownish intracellular pigment, thin-walled, very
numerous. Lamellar edge sterile. Lamellar trama inverse, up to 31 μm wide, composed of
thin-walled hyphae, 2.5–10.5 μm diam, hyaline. Pileus context undifferentiated, approx. 31
μm t ick, composed of thin-walled hyphae, 2.5–12.5 μm diam,
yaline. Pileipellis a
hymeniderm with transitions towards an epithelium and intermixed by cystidioid elements,
composed of one layer of broadly lageniform to clavate or subglobose to spheropedunculate
cells, 33–54 × 13.2–31 μm, with elongate-fusiform to filiform cystidioid elements, 70–182 ×
13.7–2 μm, t in-walled, mostly with condensed brownish intracellular pigment or rarely
with dissolved content. Caulocystidia 62–96 × 7.5–18.7 μm, elongate-clavate to elongatefusiform or filiform, thin-walled, filled with brownish content, moderately numerous. Clamp
connections absent in all parts examined.
Habit and habitat. Gregarious to disperse on decayed wood.
Specimens examined. BRAZIL. SÃO PAULO: São Paulo, Parque Estadual da
Cantareira, Núcleo Engordador, 25 Nov 2009, M. Capelari & L.A.S. Ramos MC4535
(SP417458 – HOLOTYPE).
Comments. Pluteus brunneocrinitus is morphologically close to P. crinitus,
P. necopinatus, P. seticeps (G.F. Atk.) Singer and P. thomsonii, which are also characterized
by the absence of pleurocystidia and a hymenidermal pileipellis composed of short cells
intermixed by cystidioid elements. Pluteus brunneocrinitus appears in the podospileus clade
(Fig. 1) forming a well-supported clade with P. crinitus from Brazil, P. seticeps from the
U.S.A. and an uncultured Pluteus from China (JX545226), but not related with the thomsonii
clade.
The relationship between P. brunneocrinitus and P. crinitus is very evident in both
morphological and molecular concept. Among the main morphological differences (see under
P. crinitus for full description) we can list for P. crinitus the absence of brownish fibrils at
stipe, the concolourous lamellar edges, the preponderance of subglobose basidiospores against
those globose or broadly ellipsoid, the shape, size and pigmentation of the cheilocystidia
[25–34(–49) × 12.5–21(–26) μm, colorless and short-clavate to broadly ventricose or
subglobose to spheropedunculate] and the shape and size of the caulocystidia (37–75 × 8.7–21
μm, clavate to fusoid-vesiculose).
Comparing P. brunneocrinitus and P. necopinatus, they are also morphologically
different because the latter has a less fibrillose pileus that is also distinct in color (brownishpink), concolourous lamellar edges, broader and colorless cheilocystidia, non-filiform and
shorter cystidioid elements in the pileipellis and distinct and thick-walled terminal members
of the stipitipellis hyphae (see under P. necopinatus for full description).
Capítulo V - 189
According to the description of P. seticeps presented by Minnis and Sundberg
(2010), it also differs morphologically from P. brunneocrinitus mainly by the absence of a
pileus fully covered by brownish erected fibrils with tooth-shaped scales hanging over the
edge, the presence of concolourous lamellar edge and the shape and pigmentation of the
cheilocystidia (colorless and subglobose to spheropedunculate to pyriform to clavatecylindrical), which apparently are more similar to those of P. crinitus.
Fig. 4. Pluteus brunneocrinitus (MC4535 – holotype). a. Basidiospores. b. Cheilocystidia. c. Pileipellis elements.
d. Caulocystidia. Bars = 10 µm.
Capítulo V - 190
Besides the phylogenetic position of P. brunneocrinitus and P. thomsonii (Fig. 1),
they are also separated by morphology because P. thomsonii is characterized by a blackish
and usually highly reticulate-veined pileus, the concolourous lamellar edge, the colorless
cheilocystidia with short to quite long apical projections and the pleurocystidia apparently
absent but when present they also have long apical projections (Homola 1972, Orton 1986,
Minnis and Sundberg 2010).
Pluteus cebolinhae, Menolli, Justo & Capelari, sp. nov.
FIGS. 2d, e, 5
Etymology. Named in honor of the famous Brazilian comic character ‘Cebolin a’,
(http://turmadamonica.uol.com.br/personagem/cebolinha/) whose has few strands of hair in
the head that are arranged like the sterigmata of the unusual basidia present in the lamellar
edge of the new species. Moreover, in Portuguese ‘cebolin a’ means c ive or refers to t e
diminutive of onion, and the shape of the unusual basidia and the sterigmata of P. cebolinhae
also resemble the small bulb and the short roots of chive or onion.
Pileus 20 mm diam, convex, surface brown, darker at center (N80A99M60) and paler
towards the margin (N60A99M60) to pure white alternating to the brownish tones on the
insertion points of lamellae, center veined-rugulose, margin deeply plicate-sulcate. Lamellae
free, pinkish (N10Y50M40), crowded, ventricose, ≤ 4 mm broad, with concolourous edges and
lamellulae of different lengths, 1–3 for each lamellae. Stipe 15 × 2(apex)–3(base) mm,
attenuated upwards, central, surface white, pruinose or with very small punctation overall,
with scanty basal mycelium. Odor, taste and flesh color not recorded.
Basidiospores [20/1/1] (3.7–)4.3–5.0(–5.6) × 3.7–4.3 μm [Q = (1.00–)1.16–1.35;
Qm = 1.24; Lm = 4.8 μm; Wm = 3.9 μm , broadly ellipsoid to ellipsoid, very rarely globose,
inamyloid, hyaline, smooth, thick-walled, guttulate. Basidia dimorphic: I) 16.2–23 × 7.5–10.0
μm, clavate to ventricose, frequent on lamellar sides, thin-walled, four-spored; II) 8.7–13.7 ×
8.1–10.0 μm, ovoid to balloon-shaped, present on lamellar edge, thin-walled, four-spored.
Pleurocystidia absent. Cheilocystidia 20–37 × 12.5–22 μm, short-clavate to subglobose or
spheropedunculate, colorless and hyaline, thin-walled, very rare.
Lamellar edge
heteromorphous. Lamellar trama inverse, up to 31 μm wide, composed of t in-walled
hyphae, up to 7.5 μm diam, hyaline, interwovened by oleiferous yp ae up to 2.5 μm diam.
Pileus context undifferentiated, approx. 45 μm t ick, composed of t in-walled hyphae, up to
7.5 μm diam, hyaline, interwovened by oleiferous yp ae up to 2.5 μm diam. Pileipellis an
epithelium up to 50 μm t ick, composed of one layer of subglobose to spheropedunculate
cells, 40–50 × 27–37 μm, thin-walled, filled with evenly dissolved brownish intracellular
Capítulo V - 191
pigment or almost colorless and hyaline for those near the pileus margin. Caulocystidia 22–68
× 12.5–27 μm, clavate, t in-walled, mostly colorless and hyaline or rarely with very pale
straw content. Clamp connections absent in all parts examined.
Habit and habitat. Solitary on decayed wood.
Specimen examined. BRAZIL. SÃO PAULO: São Paulo, Reserva Biológica do Alto
da Serra de Paranapiacaba, 20 Jun 2007, C.A. Canavese & F. Karstedt CAC39 (SP417455 –
HOLOTYPE).
Fig. 5. Pluteus cebolinhae (CAC39 – holotype). a. Basidiospores. b. Basidia type I. c. Basidia type II.
d. Cheilocystidia. e. Pileipellis elements. f. Caulocystidia. Bars = 10 µm.
Capítulo V - 192
Comments. Pluteus cebolinhae is characterized by a veined and deeply plicatesulcate pileus with the extreme margin colored with alternating shades of brown and white,
the dimorphic basidia, the absence of pleurocystidia, the rare cheilocystidia and the clavate
caulocystidia. Some of these characteristics are shared with P. crenulatus, a species recently
described from Dominican Republic (Justo et al. 2012) and phylogenetically close to
P. cebolinhae (Fig. 1). The deeply sulcate pileus with the margin alternating brown and
whitish shades and the absence of pleurocystidia are the main characteristics in common
between them. However, P. crenulatus has a pileus with a crenulate margin, non-dimorphic
basidia, crowded cheilocystidia forming a well-developed strip, and pileipellis elements and
caulocystidia in more varied shapes (Justo et al. 2012).
The phylogenetic position of P. crenulatus and P. cebolinhae is still unresolved.
Justo et al. (2012) showed the isolate placement of P. crenulatus as sister to all taxa in sect.
Celluloderma except of P. diettrichii, although with no statistical support. Now, P. crenulatus
and P. cebolinhae appear in both MP and ML analyses in a well-supported clade external to
all taxa in sect. Celluloderma including P. diettrichii (Fig. 1).
Among the other species of Pluteus sect. Celluloderma lacking pleurocystidia and
with a epithelial pileipellis with uniform elements, such as P. diettrichii, P. insidiosus,
P. paucicystidiatus, P. poliocnemis and P. rimulosus, P. cebolinhae differs remarkably in the
macroscopic aspect, mainly in the deeply plicate-sulcate pileus margin, and also in nondimorphic basidia and in the shape and size of the basidiospores, cheilocystidia and
caulocystidia (Kühner and Romagnesi 1956, Vellinga and Schreus 1985). Nevertheless,
P. paucistidiatus is positioned in the aurantiorugosus clade, closer to P. iguazuensis from
Brazil and P. stenotrichus from Dominican Republic, and so not phylogenetically related to
P. cebolinhae (Fig. 1).
Pluteus crinitus, Menolli & Capelari, sp. nov.
FIGS. 2g, h, 6
Etymology. The name refers to the hairy pileus surface.
Pileus 7–16 mm diam, convex then applanate, slightly depressed at center, chestnut
brown (N80Y60M30-40) and darker at center, surface fully covered by brownish erected fibrils,
densely tomentose-fibrillose to spinulose, sometimes cracking and showing the whitish flesh,
margin not striate or sulcate but with tooth-shaped scales hanging over the edge. Lamellae
free, pinkish, subdistant, ventricose, ≤ 3.5 mm broad, with concolourous edges and 1–3
lamellulae for every one series of lamellae. Stipe 12–18 × <1(apex)–1(base) mm, slightly
attenuated upwards, with a small subbulbous base, central, surface translucent-white, covered
Capítulo V - 193
by very small and dispersed whitish fibrils at base, with scanty tomentose basal mycelium.
Odor, taste and flesh color not recorded.
Basidiospores [20/1/1] (4.3–)5.0 × 4.3 μm [Q = (1.00–)1.16; Qm = 1.15; Lm = 4.9
μm; Wm = 4.3 μm , subglobose, very rarely globose, inamyloid, hyaline, smooth, thickwalled, guttulate. Basidia 21–23 × 6.2–7.5 μm, clavate to ventricose-clavate, thin-walled,
four-spored. Pleurocystidia absent. Cheilocystidia 25–34(–49) × 12.5–21(–26) μm, shortclavate to broadly ventricose or subglobose to spheropedunculate, colorless and hyaline, thinwalled, not numerous. Lamellar edge heteromorphous. Lamellar trama inverse, up to 37 μm
wide, composed of thin-walled hyphae, 2.5–12.5 μm diam, hyaline. Pileus context
undifferentiated, approx. 31 μm t ick, composed of thin-walled hyphae, 2.5–12.5 μm diam,
hyaline. Pileipellis hymeniderm with transitions towards an epithelium and intermixed by
cystidioid elements, composed of one layer of subglobose, spheropedunculate or clavate to
vesiculose cells, 32–67 × 20–29 μm, with elongate-fusiform to filiform cystidioid elements,
70–180 × 12.5–22 μm, t in-walled, mostly with evenly dissolved brownish intracellular
pigment or sometimes with few condensations. Caulocystidia 37–75 × 8.7–21 μm, clavate to
fusoid-vesiculose, thin-walled, filled with light brown content, very rare and sparse. Clamp
connections absent in all parts examined.
Habit and habitat. Gregarious to disperse on decayed wood.
Specimens examined. BRAZIL. AMAZONAS: Manaus, Instituto Nacional de
Pesquisas da Amazônia (INPA), Campus Aleixo I, 27 Jun 2011, F. Karstedt FK2064
(INPA241254 – HOLOTYPE, SP445855 – ISOTYPE).
Comments. Pluteus crinitus is also one of the few species without pleurocystidia and
having a pileipellis with cystidioid elements. Although its molecular closeness with
P. brunneocrinitus (Fig. 1), they are different in morphology based on the characteristics
discussed above (see comments under P. brunneocrinitus), such as the brownish fibrils at
stipe, the pigmented lamellar edge, the shape of the basidiospores, the shape, size and
pigmentation of the cheilocystidia and the shape and size of the caulocystidia.
Pluteus crinitus is also closely related to P. necopinatus and P. seticeps. However,
P. necopinatus has a less fibrillose and pinkish-brown pileus, slightly bigger basidiospores,
broader and differently shaped cheilocystidia, non-filiform and shorter cystidioid elements in
the pileipellis, and distinct thick-walled hyphae of the stipitipellis (see under P. necopinatus
for full description). Pluteus seticeps is also morphologically different from P. crinitus mainly
in the macroscopic appearance of the pileus that is not fully covered by brownish erected
fibrils and by the presence of dark brown fibrils all over the stipe (Minnis and Sundberg
Capítulo V - 194
2010). In addition, the sequence of P. necopinatus from Brazil and those of P. seticeps from
the U.S.A. are in separate branches from P. crinitus and P. brunneocrinitus.
Fig. 6. Pluteus crinitus (FK2064 – holotype). a. Basidiospores. b. Cheilocystidia. c. Pileipellis elements.
d. Caulocystidia. Bars = 10 µm.
Pluteus diptychocystis Singer, Sydowia 8: 123. 1954.
FIGS. 2i, 7
Pileus 22 mm diam, convex-campanulate, umbonate, dark brown to blackish
(N90A60M50) at center and greyish-brown (N80A99M50) towards the margin, covered by
Capítulo V - 195
appressed fibrils, densely tomentose-fibrillose with blackish, spinulose squamules at center,
sometimes exposing at margin the paler background between the fibrils, margin not striate or
sulcate. Lamellae free, pinkish, crowded, ventricose, with concolourous edges and lamellulae
of different lengths, 1–2 for each lamellae. Stipe 45 × 2(apex)–4(base) mm, attenuated
upwards, base sometimes clavate to subbulbous, central, surface whitish-cream, with very
small brownish punctation mainly in the mid point, with scanty basal mycelium. Odor, taste
and flesh color not recorded.
Basidiospores [60/3/3] (5.6–)6.2–7.5(8.7) × 5.0– .2 μm (Q = 1.11–1.50; Qm = 1.25;
Lm = . μm; Wm = 5.3 μm , subglobose to ellipsoid, inamyloid, hyaline, smooth, thickwalled, guttulate. Basidia 27–32 × 7.5–8.7 μm, elongate-clavate, thin-walled, four-spored.
Pleurocystidia 41–81 × 16.2−27 μm, ampullaceous, clavate or lageniform to slightly
ventricose, mostly with a median or apical outer collar adhered to the wall that detaches and
breaks easily, sometimes with basal septum, colorless and hyaline, thin-walled, sparse and
moderately abundant. Cheilocystidia 40–65 × 12.5–17.5 μm, similar to the pleurocystidia or
also vesiculose-ventricose and mostly also with the outer collar, sometimes with basal
septum, colorless and hyaline, thin-walled, abundant. Lamellar trama inverse, up to 50 μm
wide, composed of thin-walled hyphae, 2.5–15.0 μm diam, sometimes interwovened by
inflated yp ae up to 21 μm diam, hyaline. Pileus context undifferentiated, approx. 81 μm
thick, composed of thin-walled hyphae, 2.5–15.0 μm diam, yaline. Pileipellis a cutis, approx.
100 μm t ick, composed of parallel chains of cylindrical and thin-walled hyphae, individual
terminal elements 90–160 × 10.0–12.5 μm, sometimes with ascendant and suberect elements,
mostly with rounded apex or gradually attenuated towards the apex (6.0–10.0 μm broad),
sometimes with incrusted wall, filled with evenly dissolved brownish intracellular pigment.
Caulocystidia absent. Stipitipellis composed of thin-walled and yellowish hyphae, 3.7–8.7 µm
diam. Clamp connections absent in all parts examined.
Habit and habitat. Solitary on litter.
Specimens examined.
A IL. IO
ANDE DO NO TE: a a Formosa, eserva
Particular do Patrimônio Natural Mata Estrela, 14 Jul 2010, Menolli Jr. et al. NMJ184
(SP445829 . S O PA LO: Canan ia, Par ue Estadual da Il a do Cardoso, between Morro
Tr s Irm os and Ipanema, 10 Apr 1985, M. Capelari MC247 (SP193821 as P. cubensis).
Additional specimen examined. ARGENTINA. Patagonia, Parque Nacional Nahuel
Huapi, Isla Victoria, 17 May 1952, R. Singer M725 (MICH, SP – ISOTYPE).
Comments. Pluteus diptychocystis was described from Argentina (Singer 1954a) and
also recorded from Chile (Singer 1969) and Brazil (Pegler 1997). However, the
Capítulo V - 196
re-examination of the material studied (Eiten & Goodland 6313 – SP106367) by Pegler
(1997) from São Paulo State is actually P. rimosoaffinis (see description under this species).
Fig. 7. Pluteus diptychocystis (NMJ184). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia. d. Pileipellis
Morphologically, P. diptychocystis is easily recognized by a dark brown to blackish
fibrillose pileus with blackish and spinulose squamules at center and by the distinctive pleuroand cheilocystidia with an outer and fragmentary collar adhered to the wall in the median or
apical region. Similar ornamentation is also observed on the pleurocystidia of P. striatocystis
Pegler reported from Kenya (Pegler 1977) and recently from Brazil (Menolli et al. 2013b).
Capítulo V - 197
However, a distinctive character of P. striatocystis is the outer collar that is evenly striate
instead of smooth as observed in P. diptychocystis.
The molecular analyses places P. diptychocystis in the ephebeus clade as an external
branch to P. riberaltensis var. conquistensis and other two sequences of P. cf. riberaltensis
from Costa Rica, and closer to sequences of P. ephebeus (Fr.) Gillet and P. aff. ephebeus.
Although the record by Pegler (1997) is excluded, the occurrence of P. diptychocystis is
maintained in Brazil from the states of Rio Grande do Norte and São Paulo.
Pluteus cf. fastigiatus Singer, in Singer & Digilio, Lilloa 25: 259. 1952.
FIGS. 2j, 8
Pileus 52 mm diam, conic-convex, with a truncate umbo that is slightly depressed at
center, dark brown, covered by appressed and radially arranged fibrils, finally squamulose at
depressed umbo with the fibrils appearing radially split outside the central disc and sometimes
exposing at margin the paler background, margin not striate or sulcate. Lamellae free, pinkish,
crowded, ventricose, with concolourous edges and lamellulae of different lengths, 1–2 for
each lamellae. Stipe 74 × 6(apex)–8(base) mm, subequal, central, whitish-cream on
background with small brownish fibrils overall, with scanty basal mycelium. Odor, taste and
flesh color not recorded.
Basidiospores [20/1/1] (5.0–)5.6–6.2 × (4.3– 5.0 μm (Q = 1.12–1.24; Qm = 1.17;
Lm = 5.8 μm; Wm = 4.9 μm , subglobose to broadly ellipsoid, inamyloid, hyaline, smooth,
thick-walled, guttulate. Basidia 19–31 × 5.0–8.7 μm, clavate to elongate-clavate, thin-walled,
four-spored. Pleurocystidia 51–80 × 12.5−26 μm, clavate to fusiform-clavate or slightly
lageniform, sometimes with an apical mucilage, colorless and hyaline, thin-walled, sparse and
not numerous. Cheilocystidia 39–75 × (10.0–)15.0−31 μm, clavate to fusiform-clavate,
colorless and hyaline, thin-walled, numerous and organized in fascicules. Lamellar edge
sterile. Lamellar trama inverse, up to 75 μm wide, composed of thin-walled hyphae, 2.5–10.0
μm diam, hyaline. Pileus context undifferentiated, approx. 125 μm t ick, composed of thinwalled hyphae, 2.5–10.0 μm diam,
yaline. Pileipellis a cutis, approx. 100 μm t ick,
composed of parallel chains of cylindrical and thin-walled hyphae, individual terminal
elements 80–154 × 7.5–11.2 μm, mostly wit an acute apex (2.5–3.7 μm broad), filled with
evenly dissolved brownish intracellular pigment. Caulocystidia absent. Stipitipellis composed
of thin-walled hyphae, 6.2–17.5 µm diam, colorless and hyaline. Clamp connections absent in
all parts examined.
Capítulo V - 198
Fig. 8. Pluteus cf. fastigiatus (NKI12). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia. d. Pileipellis
Habit and habitat. Solitary on soil.
Specimens examined. BRAZIL. Amazonas: Instituto Nacional de Pesquisas da
Amazônia (INPA), Campus III – V-8, 25 Abr 2011, N.K. Ishikawa & R. Vargas-Isla NKI12
(INPA239970).
Comments. The collection NKI12 is undoubtedly morphologically close to
P. fastigiatus described from Argentina (Singer and Digilio 1952), mainly by the macroscopic
Capítulo V - 199
characteristics of the pileus surface, the brownish stipe, and the humicolous habitat. However,
some micromorphological differences observed between our collections and Singer’s
descriptions (Singer and Digilio 1952, Singer 1956, 1959) led us to maintain NKI12 as
inconclusive.
Following Singer (1959), some characteristics observed in P. fastigiatus are different
from those found in NKI12: i) presence of slightly larger basidiospores (6.5–7.3 × 5–6.5 µm)
and occasionally some even larger (9–11 × 7–8.5 µm); ii) occurrence of some basidia with
only one, two or three sterigmata; iii s ape of t e pleurocystidia described as “almost
exclusively ventricose-ampullaceous
(…
to
ampullaceous-subcapitate,
and
mostly
pedicellate, sometimes with spiral bodies inside but always hyaline, with thin to moderately
thickened wall without incrustation, without prongs or merely with nodose or small spinulose
e crescencies at ape , not metuloid”; iv s ape of t e c eilocystidia described as “vesiculosesubampullaceous, more rarely ventricose-ampullaceous, or ampullaceous, or vesiculoseclavate to vesiculose-submucronate”; v t e broadly rounded tips in t e pileipellis cells.
Singer (1959) placed P. albostipitatus (Dennis) Singer and P. cubensis into the same
morphological group of P. fastigiatus by the presence of long and radial fibrils in the pileus
surface and by the white to cinereous stipe with darker fibrils. However, P. albostipitatus is
currently classified in sect. Pluteus by the presence of unusual pleurocystidia with slightly
thickened to thickened wall and some small apical outgrowths (Menolli et al. 2010, 2013a,
Justo et al. 2011a, b). Based on the characteristics described by Singer (1959) for the
pleurocystidia of P. fastigiatus, maybe it could be better classified in sect. Pluteus like
P. albostipitatus and NKI12 could be described as a new species. As we were unable to
examine the type of P. fastigiatus, we prefer to maintain NKI12 as P. cf. fastigiatus, although
seems to be evident the differences between them.
The re-examination of the holotype of P. cubensis (Menolli et al. 2013b) revealed it
has larger basidiospores [7.5–8.7(–10.0) × (5.0–)6.2–7.5(–8.7 μm] and furthermore a
Brazilian collection identified as P. cubensis appears in the molecular analyses as a external
branch to the aurantiorugosus clade (Fig. 1) and so not phylogenetically related to NKI12.
Pluteus cf. fastigiatus appears in the ephebeus clade (Fig. 1) forming a wellsupported clade with other species with a cutis-like pileipellis, viz. P. fenzlii, P. mammillatus,
P. cf. fuliginosus, P. aureolatus and other two undetermined species (AJ535 and AJ478). The
presence of terminal members of the pileipellis cells with an acute to subacute apex is a
characteristic shared by NKI12, P. fuliginosus and other species classified by Singer (1959) in
the stirps fuliginosus, such as P. argentinensis Singer, P. pluvialis Singer, P. rhoadsii Murrill,
P. yungensis Singer. However all of them are different from NKI12. Pluteus argentinensis has
Capítulo V - 200
a trichohymenidermal pileipellis like the members of sect. Hispidoderma (Menolli et al.
2013c); P. fuliginosus has some pronged pleurocystidia (Smith and Stuntz 1958, Banerjee and
Sundberg 1993); P. pluvialis has a long plicate-sulcate pileus and pure white stipe (Singer
1959); P. rhoadsii has globose basidiospores and some pigmented pleurocystidia (Banerjee
and Sundberg 1993); and P. yungensis has bigger basidiospores [5.8–8.2 × 5.0–7.0(–7.5 μm]
and white stipe (Singer 1959).
Pluteus cf. fuliginosus Murrill, N. Amer. Fl. 10: 134. 1917.
FIGS. 2k, l, 9
Pileus 70 mm diam, applanate, with a distinct umbo, umbonate, surface dark brown
(N80Y99M50), densely covered by appressed and radially arranged fibrils, surface minutely
cracking around the umbo and showing the cream flesh, margin striate over at least one-third
of the radius. Lamellae free, pinkish, crowded, ventricose, ≤ 6 mm broad, with concolourous
edges and with scarce lamellulae. Stipe 92 × 6(apex)–10(base) mm, attenuated upwards,
central, surface greyish-brown (N10Y40M20), shiny and slightly longitudinally striate. Odor,
taste and flesh color not recorded.
Basidiospores [20/1/2] 6.2–6.8(7.5) × 5.0–5.6(– .2 μm [Q = (1.11–)1.21–1.24;
Qm = 1.23; Lm = .3 μm; Wm = 5.2 μm], broadly ellipsoid, rarely subglobose, inamyloid,
hyaline, smooth, thick-walled, guttulate. Basidia 25–27 × 6.2–7.5 μm, elongate-clavate to
versiform,
thin-walled,
four-spored.
Pleurocystidia
60–74
×
12.5−15.0(–22)
μm,
ampullaceous to fusoid-ventricose, usually with 2 small acute protuberances and slightly
thick-walled at apex, mostly with apical mucilage where there are a mass of basidiospores
adhered, colorless and hyaline, thin-walled, sparse and numerous. Cheilocystidia 44–66 ×
11.2–20 μm, ampullaceous to fusoid-ventricose or clavate, usually with apical mucilage
where there are a mass of basidiospores, colorless and hyaline, thin-walled, abundant.
Lamellar edge sterile. Lamellar trama inverse, up to 75 μm wide, composed of thin-walled
hyphae, 2.5–10.0 μm diam, sometimes interwovened by oleiferous yp ae up to 2.5 μm diam,
hyaline. Pileus context undifferentiated, up to 400 μm t ick, composed of t in-walled hyphae,
up to 6.2 μm diam, sometimes interwovened by oleiferous yp ae up to 2.5 μm diam, hyaline.
Pileipellis a cutis composed of parallel chains of fusoid and thin-walled hyphae, individual
terminal elements 95–115 × 16.0–20 μm, sometimes with ascendant and suberect elements,
mostly with an attenuated to subacute apex (4.0– .0 μm broad), filled with evenly dissolved
brownish intracellular pigment. Caulocystidia absent. Stipitipellis composed of thin-walled
hyphae, 3.7–15.0 µm diam, colorless and hyaline. Clamp connections absent in all parts
examined.
Capítulo V - 201
Fig. 9. Pluteus cf. fuliginosus (FK2158). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia. d. Pileipellis
Specimens examined. BRAZIL. SÃO PAULO: São Paulo, Parque Estadual das
Fontes do Ipiranga, 20 Abr 2012, F. Karstedt & M. Capelari FK2158 (SP445827).
Comments. Pluteus fuliginosus is a barely documented species known by the type
from the U.S.A. – New York (Murril 1917) with additional records from Trinidad and
Jamaica (Dennis 1953). Singer (1956) also reported P. fuliginosus from Argentina but
safeguarding the identification of the collections that after were described as P. argentinensis
Singer (Singer 1959).
Capítulo V - 202
Smith and Stuntz (1958) and Banerjee and Sundberg (1993) re-examined the
holotype of P. fuliginosus and they reported the presence of sterigma-like projections in the
pleurocystidia apex, which is most likely the main distinctive character for this species.
Dennis (1953) and Singer (1956) also re-examined the type collection but without mention of
any apical projection in the cystidia. Smith and Stuntz (1958) also reported the presence of
clamp-connections for P. fuliginosus, although the authors emphasized that they could be
considered absent for all practical purposes because only one recognizable clamp connection
was found after hours of search. We did not find clamp connection in our material and they
were not found in any other type re-examination (Dennis 1953, Singer 1956, Banerjee and
Sundberg 1993). Another characteristic observed only by Smith and Stuntz (1958) was the
presence of unusual long basidia (up to 44 μm long and also pronged c eilocystidia.
Singer (1986) considered P. fuliginosus in a distinctive stirps “c aracterized by t e
subacute or acuminate-subacute terminal cells of the epicutal trichodermium”. However, t e
organization of a trichodermal pileipellis instead of a cutis-like pileipellis as observed in our
collection is also uncertain for P. fuliginosus because according to the type re-examination
made by Smith and Stuntz (1958 , t e pileipellis was described only as aving “cuticular
elements of terminal cells in fascicules and cells elongate-fusoid, 100–220 × 10–20 μm”.
Considering the few number of preserved collections identified as P. fuliginosus and
that some incongruities about its morphology could be noted, we prefer to maintain FK2158
as inconclusive until more collections have become available from North America for an
accurate morphological and molecular comparison with the Brazilian collection. We should
not disregard the ecological differences between FK2158 and the type of P. fuliginosus,
which was originally described from a temperate/sub-boreal forest (Adirondack Mountains,
NY, U.S.A.) on a white-pine stump (Murril 1917).
According to the current molecular analyses (Fig. 1), FK2158 is positioned close to
P. fenzlii and P. mammillatus and forming a well-supported clade with P. cf. fastigiatus and
other two undetermined collections (AJ535 and AJ478) from the Dominican Republic and the
Virgin Islands.
Pluteus hispidulussimilis, Menolli, Justo & Capelari, sp. nov.
FIGS. 2m, n, 10
Etymology. The name refers to the morphological and phylogenetic relationship with
P. hispidulus.
Pileus 12 mm diam, convex, not umbonate, brownish-black at center (N90A99M30)
and greyish-brown (N80A99M35) towards the margin, surface fully covered by brownish or
greyish appressed fibrils, densely tomentose-fibrillose with blackish and spinulose squamules
Capítulo V - 203
at center, margin not striate or sulcate but slightly fringed with some long whitish fibrils.
Lamellae free, pinkish, crowded, with concolourous edges and lamellulae for every 1–2
lamellae. Stipe 24 × 1(apex)–2(base) mm, slightly attenuated upwards, with a small
subbulbous base, central, surface white and entirely covered by whitish fibrils mainly at base,
slightly longitudinally striate at apex, with scanty tomentose basal mycelium. Odor, taste and
Fig. 10. Pluteus hispidulussimilis (NMJ179 – holotype). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia.
d. Pileipellis elements. Bars = 10 µm.
Basidiospores [20/1/1] 5.0–5.6 × (4.3–)5.0(–5.
Capítulo V - 204
μm [Q = 1.00–1.12(–1.16);
Qm = 1.04; Lm = 5.2 μm; Wm = 5.0 μm], globose to subglobose, very rarely broadly
ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate. Basidia 20–26 × 7.5–8.7 μm,
clavate, thin-walled, four-spored. Pleurocystidia 27–41 × 7.5−17.5 μm, ventricose to
lageniform or elongate-clavate, colorless and hyaline, thin-walled, very rare or apparently
absent in some slices. Cheilocystidia 29–64 × 8.7–21 μm, very variable in shape, clavate,
ampullaceous, fusiform, capitate or spheropedunculate, colorless and hyaline, thin-walled,
moderately numerous. Lamellar edge heteromorphous. Lamellar trama inverse, up to 31 μm
wide, composed of thin-walled hyphae, 2.5–10.0 μm diam, hyaline. Pileus context
undifferentiated, approx. 81 μm t ick, composed of thin-walled hyphae, 2.5–10.0 μm diam,
hyaline. Pileipellis a cutis, approx. 100 μm t ick, composed of parallel chains of cylindrical
and thin-walled hyphae, with short individual terminal elements 42–76 × 12–18 μm,
sometimes with ascendant and suberect elements mainly at pileus center, mostly with rounded
apex or gradually attenuated towards the apex (6.0–10.0 μm broad), sometimes with incrusted
wall at apex, filled with evenly dissolved brown intracellular pigment. Caulocystidia absent.
Stipitipellis composed of thin-walled hyphae, 2.5–11.2 µm diam, colorless and hyaline.
Specimens examined.
A IL. IO
ANDE DO NO TE: a a Formosa, Reserva
Particular do Patrimônio Natural Mata Estrela, 14 Jul 2010, Menolli Jr. et al. NMJ179
(SP417459 – HOLOTYPE).
Comments. Pluteus hispidulussimilis is characterized by a blackish and densely
tomentose-fibrillose pileus, the globose to subglobose basidiospores, the rarely present
pleurocystidia and the very versiform cheilocystidia.
The current molecular analyses (Fig. 1) showed the phylogenetic relationship
between P. hispidulussimilis and other species with absent to very rare pleurocystidia and a
cutis-like pileipellis, viz. P. hispidulus and P. karstedtianus.
Pluteus hispidulus is a barely recorded species that seems to be restricted to Europe
(Kühner and Romagnesi 1956, Orton 1986, Justo and Castro 2007) and is morphologically
different from P. hispidulussimilis by the non-globose basidiospores, and the mostly clavate
cheilocystidia (Kühner and Romagnesi 1956, Vellinga and Schreus 1985). Vellinga and
Schreurs (1985) presented P. hispidulus var. cephalocystis Schreurs as different from the type
variety by the presence of subcapitate to capitate narrowly utriform to narrowly clavate
cheilocystidia and by the apparently almost globose basidiospores as illustrated by them.
Orton (1986) also reported for one collection of P. hispidulus more versiform cheilocystidia,
Capítulo V - 205
varying from clavate or fusiform to lageniform, utriform or capitate lageniform, but
apparently it represents a third taxon with transitional characteristics between P. hispidulus
var. hispidulus and P. hispidulus var. cephalocystis and more related morphologically to
P. hispidulussimilis.
Delimitation of P. hispidulus involves also the morphological relationship with
P. exiguus (Pat.) Sacc., another very rarely recorded species in Europe (K
ner and
Romagnesi 1956, Vellinga and Schreurs 1985, Justo and Castro 2007), which is characterized
by the clavate cheilocystidia with a distinct narrow apex or apical projections and by t e
palisade-like to tric odermal pileipellis (K
ner and omagnesi 1956, Vellinga and Schreurs
1985, Orton 1986) that is characteristic of species currently classified in sect. Hispidoderma
(Justo et al. 2011a, b). Even that the morphological differences between P. hispidulus and
P. hispidulussimilis are evident, the identification of some sequences used in the molecular
analyses require further attention because there are sequences identified as P. hispidulus
(ARAN8200509 and A1882) in different clades and one of these (A1882) clustered with a
P. exiguus sequence (FJ774083), but this question falls outside the scope of this paper.
Menolli et al. (2013c) include in the molecular analyses of sect. Hispidoderma a sequence of
P. cf. exiguus from Spain that most likely corresponds to the true P. exiguus.
Pluteus karstedtianus differs from P. hispidulussimilis in the chestnut but not dark or
blackish pileus, the sulcate striate pileus margin, the translucent and fewer fibrillose stipe and
non-versiform and rarely present to almost absent cheilocystidia (see under P. karstedtianus
for full description).
Pluteus iguazuensis Singer, Trans. Brit. Mycol. Soc. 39: 201. 1956.
FIG. 11
Pileus 21 mm diam, convex then applanate, dark brown on disc and paler towards
the margin, smooth at center and sulcate-striate over half of the radius. Lamellae free, pinkish,
crowded, ventricose, with concolourous edges and few lamellulae. Stipe 37 × 1.5(apex)
–2(base) mm, almost equal, central, surface whitish-cream with brownish base, longitudinally
striate, with scanty basal mycelium. Odor, taste and flesh color not recorded.
Basidiospores [20/1/1] (5.0–)5.6–6.2 × 5.0–5.6(–6.2) µm (Q = 1.00–1.12; Qm =
1.06; Lm = 5.8 µm; Wm = 5.5 µm), globose to subglobose, inamyloid, hyaline, smooth, thickwalled, guttulate. Basidia 27–34 × 7.5–10.0 µm, elongate-clavate, thin-walled, four-spored.
Pleurocystidia 41–59(–71) × (14.7–)21−29(–36) μm, clavate to ventricose-vesiculose,
colorless and hyaline, thin-walled, sparse and not numerous. Cheilocystidia 26–39 × 8.7–20
μm, preponderantly clavate, colorless and hyaline, thin-walled, moderately numerous.
Lamellar edge fertile. Lamellar trama inverse, approx. 31 µm wide, composed of thin-walled
Capítulo V - 206
hyphae, 2.5–11.2 µm diam, interwovened by oleiferous yp ae up to 5.0 μm diam, hyaline.
Pileus context undifferentiated, up to 37 µm thick, composed predominantly of oleiferous
hyphae up to 5.0 µm diam and intermixed by thin-walled and inflated hyphae, 12.5–18.7 µm
diam, hyaline. Pileipellis an epithelium up to 50 μm t ick, composed of one layer of
subglobose, spheropedunculate or clavate cells, 28–50 × 16–31 μm, thin-walled, filled with
evenly dissolved brownish intracellular pigment or rarely slightly few condensations.
Caulocystidia apparently absent. Stipitipellis composed of thin-walled hyphae, colorless and
hyaline but with elongated terminal members filled with brownish content, 56–77 × 8.7–11.2
μm. Clamp connections absent in all parts examined.
Specimen examined. BRAZIL. Amazonas: Instituto Nacional de Pesquisas da
Amazônia (INPA), Campus III – V-8, 25 Apr 2011, N.K. Ishikawa & R. Vargas-Isla NKI10
(INPA239968).
Fig. 11. Pluteus iguazuensis (NKI10). a. Basidiospores. b. Pleurocystidia. c. Cheilocystidia. d. Pileipellis
Capítulo V - 207
Comments. Pluteus iguazuensis was described from Argentina (Singer 1956) and
also recorded from Brazil by Meijer (2006 as P. cf. iguazuensis) and Drechsler-Santos et al.
(2007). However, the re-examination of the material studied by Meijer (2006) was
re-identified as P. crassus by Menolli et al. (2013b). About the record of Drechsler-Santos et
al. (2007), we were unable to certify the identification because the author only present the
species in a list and without indication of the collection studied.
Morphologically P. iguazuensis is recognized by a sulcate and non-venose brownish
pileus, the globose to subglobose basidiospores and by the similarity in the shape of the
clavate to ventricose-vesiculose pleuro- and cheilocystidia. Singer (1959) placed P. fallax and
P. tucumanus Singer into the same morphological group (stirps tucumanus) that
P. iguazuensis. However, according to Singer (1959), P. fallax has a rimose but not distinctly
sulcate pileus and P. tucumanus has an appendiculate-crenate pileus margin and smaller
basidiospores (3.8–5.5 × 2.7–3.8 µm).
The molecular analyses showed P. iguazuensis positioned in the aurantiorugosus
clade as an external branch to all yellow-orange species of this clade and clustered in a wellsupported clade with P. paucicystidiatus, P. stenotrichus and a undetermined collection
(AJ842) from the Dominican Republic (Fig. 1). This work confirms the occurrence of
P. iguazuensis from Brazil and represents the first record from the Amazonian region.
Pluteus karstedtianus, Menolli, Justo & Capelari, sp. nov.
FIGS. 2o, p, 12
Etymology. The name is in honor to Fernanda Karstedt by her contribution for more
than forty specimens of Pluteus collected between 2006 and 2012 around Brazil. F. Karstedt
is the collector of almost 12% of all Brazilian materials of Pluteus studied by the first author
since 2007. Moreover the collection FK637 of P. karstedtianus is probably one of the first
Pluteus specimens collected by her.
Pileus 13–22 mm diam, convex to plane-convex, sometimes subumbonate and
slightly depressed around the umbo, chestnut-brow to coffee-brown, darker at center and paler
towards the margin, surface fibrillose-squamulose, fully covered by brownish fibrils,
punctate-squamulose to spinulose at center with the fibrils appearing split outside the central
disc and sometimes concentrating on the insertion points of lamellae, alternating to the
brownish and pale cream shades, margin sulcate-striate over half of the radius, sometimes
exposing the whitish background, with some long brownish fibrils at margin. Lamellae free,
cream to pinkish, close but not crowded, with concolourous edges and short lamellulae for
every one series of lamellae. Stipe 15–23 × 1–1.5(apex)–1.5–2.0(base) mm, slightly
attenuated upwards, central, translucent to greyish-white, covered by very small whitish
Capítulo V - 208
fibrils at base, with scanty tomentose basal mycelium. Odor, taste and flesh color not
recorded.
Basidiospores [40/2/2] 5.0–5.6(–6.2) × 4.3–5. μm [Q = 1.00–1.12(–1.16); Qm =
1.05; Lm = 5.2 μm; Wm = 5.0 μm , globose to subglobose, very rarely broadly ellipsoid,
inamyloid, hyaline, smooth, thick-walled, guttulate. Basidia 20–27 × 7.5–8.7 μm, clavate to
ventricose, thin-walled, four-spored. Pleurocystidia absent. Cheilocystidia 26–37 × 7.5–13.7
μm, fusoid-ventricose to slightly lageniform, very rare or apparently absent in some slices.
Lamellar edge heteromorphous. Lamellar trama inverse, up to 37 μm wide, composed of
thin-walled hyphae, 2.5–8.7 μm diam, hyaline. Pileus context undifferentiated, approx. 42 μm
thick, composed of thin-walled hyphae, 1.2–6.2 μm diam, sometimes interwovened by
inflated hyphae up to 20 μm diam, hyaline. Pileipellis a cutis, approx. 100 μm t ick,
composed of parallel chains of cylindrical and thin-walled hyphae, with short individual
terminal elements 48–92 × 7.5–12.5 μm, sometimes with ascendant and suberect elements
mainly at pileus center, mostly with rounded apex or gradually attenuated towards the apex
(4.0–7.0 μm broad), filled with evenly dissolved brown intracellular pigment. Caulocystidia
absent. Stipitipellis composed of thin-walled hyphae, 2.5–7.5 µm diam, colorless and hyaline.
Habit and habitat. Solitary on decayed unidentified wood or on decayed caudex of
Dicksonia sp.
Specimens examined.
Serra dos
A IL.
IO DE ANEI O:
uapimirim, Par ue Nacional
rg os, Sede Guapimirim, Po o da Ponte Vel a, 6 Jan 2011, F Karstedt et al.
FK1700 (SP445828 – HOLOTYPE); SÃO PAULO: São Paulo, Parque Estadual da
Cantareira, Núcleo Engordador, 27 Apr 2006, F. Karstedt & M. Capelari FK637 (SP417456).
Fig. 12. Pluteus karstedtianus (FK1700 – holotype). a. Basidiospores. b. Cheilocystidia. c. Pileipellis elements.
Bars = 10 µm.
Capítulo V - 209
Comments. Pluteus karstedtianus is characterized by a chestnut-brow and fibrillosesquamulose pileus, the globose to subglobose basidiospores, the absence of pleurocystidia and
by the rarely present to almost absent cheilocystidia.
Pluteus karstedtianus is phylogenetically (Fig. 1) and morphologically close to
P. hispidulus and P. hispidulussimilis, two species also characterized by the absence or the
rare presence of pleurocystidia and a cutis-like pileipellis. However, P. hispidulus differs in
having non-globose basidiospores and clavate and more numerous cheilocystidia (Kühner and
Romagnesi 1956, Vellinga and Schreurs 1985). Nevertheless, P. karstedtianus fell in a
separate branch to the European sequences identified as P. hispidulus (Fig. 1). Pluteus
hispidulussimilis also differs from P. karstedtianus in the blackish pileus with non-sulcate
margin and the numerous and very versiform cheilocystidia (see P. hispidulussimilis for
complete description).
Pluteus necopinatus, Menolli & Capelari, sp. nov.
FIGS. 2q, r, 13
Etymology. necopinatus means ‘‘unexpected, surprising’’, in reference to t e fact
that the type material was found growing attached to a cement wall.
Pileus 14 mm diam, convex to applanate, depressed at center, pinkish-brown
(N40A20M30) and distinctly brownish at center (N50A60M50), surface fully covered by small
brownish fibrils, center entire and densely punctate-fibrillose, with the fibrils dissociating
outside the central disc and exposing the pinkish background, finely punctate-fibrillose
overall, margin slightly sulcate-striate with few short scales hanging over the edge. Lamellae
free, pinkish (N20A30M30 , subdistant, ventricose, ≤ 3.0 mm broad, with concolourous edges
and 1–2 lamellulae for every one series of lamellae. Stipe 12 × 1(apex)–1.2(base) mm,
slightly attenuated upwards, with a small subbulbous base, central, surface translucent-pink
(N10A20M10), covered by very small and dispersed brownish fibrils over all. Odor, taste and
Basidiospores [20/1/1] 5.0–5.6 × 5.0–5. μm (Q = 1.00–1.12; Qm = 1.05; Lm = 5.5
μm; Wm = 5.2 μm , globose to subglobose, inamyloid,
yaline, smoot , t ick-walled,
guttulate. Basidia 16.2–21 × 6.2–8.7 μm, clavate to ventricose-clavate, thin-walled, fourspored. Pleurocystidia absent. Cheilocystidia 37–74 × 12.5–30 μm, clavate to ventricoseclavate or almost lageniform, colorless and hyaline, thin-walled, moderately abundant.
Lamellar edge sterile. Lamellar trama inverse, up to 25 μm wide, composed of t in-walled
hyphae, 1.2–3.7 μm diam, interwovened by oleiferous yp ae up to 3.7 μm diam, yaline.
Pileus context undifferentiated, appro . 31 μm t ick, composed mostly by oleiferous yp ae
up to 3.7 μm diam, wit
thin-walled hyphae, 1.2–3.7 μm diam,
yaline. Pileipellis
Capítulo V - 210
hymeniderm with transitions towards an epithelium and intermixed by cystidioid elements,
composed of one layer of subglobose, spheropedunculate or clavate to fusoid-vesiculose cells,
39–65 × 24–32 μm, with fusoid to elongate-fusiform cystidioid elements, 72–125 × 20–37
μm, thin-walled, mostly with evenly concentrated brownish intracellular pigment or
Fig. 13. Pluteus necopinatus (FK1701 – holotype). a. Basidiospores. b. Cheilocystidia. c. Pileipellis elements.
d. Terminal elements in the stipitipellis. Bars = 10 µm.
Capítulo V - 211
sometimes almost colorless. Caulocystidia apparently absent but with some distinct terminal
members of the stipitipellis organized in fascicules of irregular and slightly thick- to thickwalled hyphae, 28–50 × 3.7–8.7 μm, filled with evenly dissolved brownish intracellular
pigment and sometimes with some points of incrusted wall. Clamp connections absent in all
parts examined.
Habit and habitat. Solitary on a cement wall.
Specimens examined. BRAZIL. RIO DE JANEIRO: Teresópolis, Parque Nacional
Serra dos
rg os, Sede Teresópolis, 6 Jan 2011, F Karstedt et al. FK1701 (SP445830 –
HOLOTYPE).
Comments. Pluteus necopinatus is closely related to other few species characterized
by the absence of pleurocystidia and a hymenidermal pileipellis composed of short cells
intermixed by cystidioid elements, viz. P. brunneocrinitus, P. crinitus, P. seticeps and
P. thomsonii. Pluteus necopinatus appears in the podospileus clade linked with no statistical
support to the clade composed of P. seticeps, P. crinitus, P. brunneocrinitus and an
uncultured Pluteus from China (JX545226), but not related with the thomsonii clade (Fig. 1).
Pluteus brunneopictus and P. crinitus are morphologically different from
P. necopinatus by all characteristics previously discussed (see discussion under these
species). Pluteus necopinatus is also distinguished from P. seticeps in the pileus color (dark
brown), the shorter and differently shaped cheilocystidia (18–50 × 11–35 μm, subglobose to
spheropedunculate to pyriform to clavate-cylindrical) and the absence of distinct thick-walled
terminal members in the stipitipellis (Minnis and Sundberg 2010).
The closest taxon of P. necopinatus is an undescribed material from the U.S.A.
(AJ838, collected in New Haven, Connecticut), which is 96.9 % similar in the ITS sequence.
Morphologically AJ838 differs in the presence of pleurocystidia, differently shaped and more
variable pileipellis elements and longer terminal elements in the stipitipellis (A. Justo,
unpublished data).
Pluteus paucicystidiatus, Menolli, Justo & Capelari, sp. nov.
FIGS. 14a–e, 15
Etymology. The name refers to the almost complete absence of cystidia.
Pileus 10–20 mm diam, shallowly and broadly obtusely conical to broadly convex
then applanate, dark brown (6F6-8) on disc, slightly paler towards the margin and sometimes
splitting, rugose to venose at center, margin brown to light brown (6E7-8) and sulcate-striate
over at least one-third of the radius. Lamellae free, white then pinkish grayish orange
(6A-B3), close to crowded, with concolourous edges and 1–3 lamellulae for every one series
Capítulo V - 212
of lamellae. Stipe 20–30 × 1.5–2 mm, almost equal, central, white to translucent, finely
pruinose. Odor, taste and flesh color not recorded.
Basidiospores [20/1/1] 6.2–7.5 × 5.0–6.2 µm (Q = 1.21–1.24; Qm = 1.22; Lm = 6.9
µm; Wm = 5.7 µm), broadly ellipsoid, inamyloid, hyaline, smooth, thick-walled, guttulate.
Basidia 17.5–24 × 6.2 µm, versiform, thin-walled, four-spored. Pleurocystidia almost absent,
it was possible to see only a single clavate element (44 × 15.0 µm) in one of many slices.
Cheilocystidia 31–45 × 10.0–25 μm, clavate to clavate-fusoid or slightly lageniform to almost
utriform, colorless and hyaline, thin-walled, rare to almost absent in some sections. Lamellar
edge subheteromorphous with few basidia and many basidioles, but not differentiated in
cystidioid elements. Lamellar trama inverse, approx. 12.5 µm wide, composed of thin-walled
hyphae, 3.7–8.7(–11.2) µm diam, interwovened by oleiferous yp ae up to 5.0 μm diam,
hyaline. Pileus context undifferentiated, up to 75 µm thick, composed predominately of
oleiferous hyphae up to 6.2 µm diam and intermixed by thin-walled hyphae, 3.7–8.7 µm
diam, hyaline. Pileipellis a epithelium with transitional elements to a hymeniderm, up to 50
μm t ick, individual elements (23–)30–50 × 10.0–37 μm, mostly spheropedunculate but few
like the cheilocystidia, clavate to clavate-fusoid or almost utriform, filled with dissolved or
sometimes condensed brownish intracellular pigment. Caulocystidia 22–68 × 12.5–27 μm,
clavate or broadly lageniform, thin-walled, filled with condensed or dissolved very pale straw
content. Clamp connections absent in all parts examined.
Habit and habitat. Solitary to subcespitose in clusters of two-three basidiomata on
rotten hardwood logs.
Specimen examined. BRAZIL. SÃO PAULO: Iporanga, Parque Estadual Turístico
do Alto Ribeira, Núcleo Ouro Grosso, 21 Mar 2007, C. Puccinelli et al. CP218 (SP394383 –
HOLOTYPE).
Comments. Pluteus paucicystidiatus is characterized by the almost complete absence
of cystidia and by the pileus rugose to venose at center with a sulcate-striate margin.
According to Singer’s (198 ) classification for sect. Celluloderma and considering the
macroscopic characteristics of P. paucicystidiatus, it could be compared to some species of
stirps longistriatus
including P. longistriatus
(Peck) Peck,
P.
neophlebophorus,
P. sanctixaverii Singer, P. variipes and P. oligocystis. However, all of these taxa, including
P. neophlebophorus and P. sanctixaverii that are most similar with P. paucicystidiatus due to
the rugose-venose pileus, have a pileipellis intermixed by cystidioid elements and presence of
cystidia (Singer 1959). Nevertheless, P. longistriatus is now accepted in sect. Hispidoderma
(Justo et al. 2011a, b, Menolli et al. 2013 a, c).
Capítulo V - 213
Fig. 14. a–e. Pluteus paucicystidiatus (CP218 – holotype). a. Basidiomata. b. Lamellar section showing the
absence of cystidia. c. Lamellar section showing the only pleurocystidia seen (black arrow). d. Pleurocystidia.
e. Cheilocystidia. f. P. rimosoaffinis (MC4607). Bars: a, f = 1 cm; b–c = 50 µm; d–e = 10 µm.
Among the species of sect. Celluloderma characterized by the absent to very rare
pleurocystidia and a hymenidermal pileipellis composed of short cells and without cystidioid
elements, P. paucicystidiatus could be compared to P. crenulatus, P. diettrichii, P. insidiosus
Vellinga & Schreurs, P. poliocnemis Kühner, P. cebolinhae and P. rimulosus Kühner &
Romagn.
Pluteus crenulatus is clearly different from P. paucicystidiatus by the deeply sulcate
pileus with a crenulate margin and the globose basidiospores. In addition, the phylogenetic
analyses (Fig. 1) put P. crenulatus closer to P. cebolinhae in an external branch to all taxa in
sect. Celluloderma. Pluteus rimulosus was considered a synonym of P. diettrichii by Vellinga
Capítulo V - 214
and Schreurs (1985) and both, together with P. poliocnemis, have longer basidiospores that
are ellipsoid to oblong. Besides, they are commonly found on soil and are apparently
restricted to Europe (Kühner and Romagnesi 1956, Vellinga and Schreurs 1985, Orton 1986,
Justo and Castro 2007). Pluteus insidiosus probably is the species morphologically closer to
Fig. 15. Pluteus paucicystidiatus (CP218 – holotype). a. Basidiospores. b. Pleurocystidium. c. Cheilocystidia.
d. Pileipellis elements. e. Caulocystidia. Bars = 10 µm.
Capítulo V - 215
P. paucicystidiatus due to the pileus color and size and shape of the basidiospores and
pileipellis cells (Vellinga and Schreurs 1985). However, according to Vellinga and Schreurs
(1985), P. insidiosus has characteristic cheilocystidia with long apical projections and, in
addition, the pileus is not distinctly sulcate neither rugose-venose as in P. paucicystidiatus.
Finally, P. cebolinhae has a deeply plicate-sulcate pileus and dimorphic basidia that differ
from P. paucicystidiatus, and there is no phylogenetic relationship between them (Fig. 1).
In the previous phylogenetic analyses of Justo et al. (2011b), P. paucicystidiatus (as
Pluteus sp. IV SP394383) was included in the romelli/aurantiorugosus clade as sister to all
yellow-orange species of this clade, and also separated from other species without cystidia
and pileipellis without cystidioid elements, such as P. diettrichii. This position was confirmed
in the analysis of Justo et al. (2012) that showed its relationship with P. stenotrichus, and
now, based on the current analyses (Fig. 1), we also show a phylogenetic relationship with
P. iguazuensis from Amazonas, Brazil.
Pluteus rimosoaffinis Singer, Trans. Brit. Mycol. Soc. 39: 211. 1956.
FIGS. 14f, 15
Pileus 25–35 mm diam, plane-convex, slightly depressed around a broad and low
umbo, dark brown (A90M90C90) at center and light brown (A80M70C70) at margin,
subvelutinous, surface cracking overall and showing the cream flesh except at umbo, center
almost entire, minutely rugose to venose, margin indistinctly sulcate. Lamellae free to
subfree, pinkish, with concolourous edges and lamellulae every 1–2 series of lamellae, under
magnifying glass it is possible to see numerous cystidia on lamellar face that are apparently
pigmented due to the pale melleous mass of basidiospores adhered to the apical mucilage
(seen under microscopy). Stipe 25–30 × 3(apex)–4–5(base) mm, tapering towards the apex,
base subbulbous, central to slightly eccentric, pale cream to slightly brownish, finely pruinose
and with finely brownish fibrils at base, slightly longitudinally striate, with scanty basal
mycelium. Odor, taste and flesh color not recorded.
Basidiospores [100/5/5] 5.0–7.5 × 5.0–6.8(–7.5) µm (Q = 1.00–1.24; Qm = 1.10; Lm
= 6.5 µm; Wm = 6.0 µm), globose to broadly ellipsoid, inamyloid, hyaline and colorless to
pale melleous, smooth, thick-walled, guttulate. Basidia 25–34 × 7.5–8.7(–10.0) µm, clavate to
versiform, thin-walled, four-spored. Pleurocystidia (42–)55–79 × 14.0–24(–29) µm, slightly
ventricose to lageniform or almost ampullaceous, sometimes fusiform-clavate or clavate,
frequently with basidiospores adhered to an apical mucilage that under magnifying glass are
visible as light brown points in the lamellar face, colorless or sometimes with very pale straw
internal dissolved content, thin-walled, sparse and moderately abundant. Cheilocystidia (32–)
45–75(–85) × (10.0–)13.7–25(–37) µm, like the pleurocystidia, with pale melleous internal
Capítulo V - 216
dissolved content or in some collections hyaline and colorless, moderately numerous.
Lamellar edge apparently sterile. Lamellar trama inverse, 50–100 µm wide, composed of
thin-walled hyphae, 6.2–12.5 µm diam, interwovened by oleiferous hyphae up to 5.0 µm
diam, hyaline. Pileus context undifferentiated, approx. 360 µm or up to 800 µm thick at pileus
center, composed of thin-walled hyphae, 5.0–12.5 µm diam, interwovened by oleiferous
hyphae up to 3.7 µm diam hyaline. Pileipellis an epithelium up to 50 μm t ick, composed of
one layer of subglobose to vesiculose cells, 30–62 × 18.7–37(–5
μm, thin-walled, filled with
evenly dissolved or condensed brownish intracellular pigment. Caulocystidia apparently
absent. Stipitipellis composed of thin-walled hyphae, colorless and hyaline but with elongated
terminal elements filled with brownish content, 52–80 × 7.5–11.2 μm. Clamp connections
absent in all parts examined.
Habit and habitat. Solitary or gregarious on decayed wood.
Specimens examined. BRAZIL. BAHIA: Itabuna, 6 Jun 1978, R. Singer B 11111
(INPA82432); RIO GRANDE DO SUL: Passo Fundo, Jaboticabal, 23 Oct. 2003, M.S. Rother
& B.M.A. Severo s.n. (RSPF327 as P. fallax); SÃO PAULO: Itapecerica da Serra, 25 Jun.
1965, Eiten & Goodland 6313 (SP106867 as P. diptychocystis); São Paulo, Parque Estadual
da Cantareira, Núcleo Engordador, 7 Nov 2011, M. Capelari & J.J.S. Oliveira MC4607
(SP416740); Parque Estadual das Fontes do Ipiranga, 19 Aug 2009, A.M. Gugliotta et al.
PEFI07/2009 (SP394379); Mogi Guaçu, Reserva Biológica de Mogi Guaçu, Fazenda
Campininha, 29-30 Jan. 1987, D.N. Pegler et al. 3896 (SP214395).
Comments. Pluteus rimosoaffinis was described from Argentina (Singer 1956) and
since then it was recorded from Brazil by Pegler (1997) and Meijer (2006 as
P. cf. rimosoaffinis) but both without complete description.
Singer (1956) mentioned in the protologue some characteristics that were not
observed in the Brazilian collections, such as the entirely white stipe, the grayish-pallid
lamellar edge, not geometrically globose basidiospores, and the presence of obpiriform
pleuro- and cheilocystidia. Moreover, we also observed some weakly pigmented
pleurocystidia.
In the Brazilian material collected and identified by R. Singer (R. Singer B 11111,
data not published) it was not possible to observe basidiospores neither cystidia to confirm
these variations. On the other hand, re-examining the collection D.N. Pegler et al. 3896, we
verified it has slightly ventricose to clavate pleuro- and cheilocystidia and broadly ellipsoid
basidiospores, instead of obpiriform cystidia and subglobose basidiospores as published by
Pegler (1997). The collections studied by Meijer (2006) and published as P. cf. rimosoaffinis
actually represent other species (Table 1, Menolli et al. 2013b).
Capítulo V - 217
Fig. 15. Pluteus rimosoaffinis (A.M. Gugliotta et al. PEFI07/2009). a. Basidiomata b. Basidiospores. c. Basidia.
d. Pleurocystidia. c. Cheilocystidia. d. Pileipellis elements. e. Caulocystidia. Bars: a = 1 cm; b–f = 10 µm.
Although Singer (1956, 1959) emphasized the obpiriform shape of the cystidia, he
also illustrated pleuro- and cheilocystidia fusiform and pedicellate to ventricose like those
found in the Brazilian collections herein examined, even characterizing the cheilocystidia as
more versiform than the pleurocystidia. Despite these differences between the Brazilian and
the Argentinean collections, up to now we accept P. rimosoaffinis with these variations and
confirm its occurrence in Brazil. The re-examination of the Argentinean collections studied
by Singer (1956, 1959) could be useful, but the holotype was not available for loan and the
Capítulo V - 218
ot er collections (including t e paratype mentioned in Singer’s papers were not found in
LIL.
Justo et al. (2011a, b) recorded as Pluteus sp. III the collection PEFI07/2009 and
confirmed its position in the cinereofuscus clade as a sister branch to Pluteus sp. II (Justo et
al. 2011a, b) that was identified as P. sapiicola by Menolli et al. (2013b). This position was
here confirmed and it was included an additional collection of P. rimosoaffinis (MC4607) in
the current analyses (Fig. 1).
ACKNOWLEDGMENTS
The authors thank the curators of INPA, RSPF and SP for the loan of the specimens studied; Noemia Kazue
Ishikawa and Iuri Goulart Baseia for helping and all assistance during the expeditions to Manaus and Natal,
respectively; N.K. Ishikawa and Iuri Goulart Baseia, as well as Fernanda Karstedt, Carla Puccinelli, Adriana
M. Gugliotta and Dennis E. Desjardin, are also thanked for photographs and/or collecting some specimens used
in this study; Flávia Ribeiro Santos for helping with DNA extraction of some samples; Klei R. Sousa for inking
the line drawings; Francisco Kuhar and Tarciso S. Filgueiras for helping with the suggestions and composition
of the new names. N. Menolli r. and M. Capelari t ank “Funda o de Amparo à Pes uisa do Estado de S o
Paulo” (FAPESP grants 04/04319-2, 09/53272-2 , “Coordena o de Aperfei oamento de Pessoal de N vel
Superior” (CAPES - PNADB grant 23038.000043/2010-43 and t e “Consel o Nacional de Desenvolvimento
Cient fico e Tecnológico” (CNP fellows ips for financial support. A. Justo acknowledges support from the
National Science Foundation grant DEB 0933081.
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Discussão Geral - 222
DISCUSSÃO GERAL
Este trabalho possibilitou ampliar e atualizar o conhecimento das espécies de Pluteus
no Brasil, com o reconhecimento de 48 espécies, incluindo nove novos registros e 11 espécies
novas, distribuídas em 11 estados brasileiros (Amazonas, Bahia, Minas Gerais, Pará, Paraná,
Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, Santa Catarina e São
Paulo).
A partir do levantamento das espécies de Pluteus já registadas para o Brasil, bem
como da realização de novas coletas, foi possível chegar ao número de 333 exsicatas
depositadas em herbários nacionais e internacionais (APÊNDICE C). A revisão de 50
exsicatas não foi possível pois os materiais não foram localizados nos herbários de origem ou
estavam impossibilitados para empréstimo. Das 283 coleções restantes, 168 foram
identificadas, correspondendo à 43 das 48 espécies consideradas neste trabalho. As demais
coleções estão em condições insuficientes para estudo ou representam espécies de outros
gêneros (APÊNDICE C). A revisão das coleções brasileiras de P. amazonicus, P. aquosus,
P. fallax e P. paraensis não foi possível pois os materiais não foram localizados ou estavam
mal preservados. Mesmo assim, essas espécies foram consideradas como ocorrentes no Brasil
e foram incluídas na chave de identificação devido à descrição diagnóstica apresentada pelos
autores (Singer 1954, 1973, 1989, Wartchow et al. 2006).
Das 48 espécies consideradas (APÊNDICE B), a seção Celluloderma é a que tem
maior representatividade, com 32 espécies, incluindo dez espécies novas para a ciência
(capítulos III e V) e cinco novos registros para o país (capítulo III). A seção Hispidoderma
está representada por nove espécies, incluindo quatro novos registros e uma espécie nova
(capítulos III e IV), enquanto que a seção Pluteus está representada por sete espécies, sendo
todas elas previamente publicadas para o país e apenas revisadas neste trabalho (capítulos I, II
e III).
A obtenção de sequências de ITS foi possível para 26 das 48 espécies, sendo que oito
das 11 espécies novas são propostas com suporte molecular (capítulo V), possibilitando a
inserção de sequências de materiais tropicais na filogenia global do gênero e contribuindo
para estabelecer relações entre espécimes geograficamente separados. Além disso, sequências
de P. cf. fastigiatus, P. cf. fernandezianus e P. cf. fuliginosus também foram incluídas nas
árvores geradas para as seções Celluloderma e Hispidoderma (capítulo IV e V), porém, novas
coletas serão necessárias para a correta identificação desses materiais.
Os estudos filogenéticos com as espécies de Pluteus permitiram uma maior
compreensão da classificação infragenérica proposta para o gênero, permitindo o
Discussão Geral - 223
reconhecimento de clados bem suportados dentro das seções Celluloderma e Hispidoderma.
Uma maior amostragem das espécies incluídas nas análises da seção Celluloderma
possibilitou melhor embasamento para considerar como artificial o uso de caracteres
morfológicos que apareceram independentemente na escala evolutiva do gênero, como, por
exemplo, as diferentes formas de superfície pilear observadas nessa seção: indiferenciada
(= seção Villosi); himeniforme ou epitelial sem elementos cistidioides (= subseção
Eucellulodermini); ou himeniforme ou epitelial com elementos cistidioides (= subseção
Mixtini).
Estudos multigênicos são indicados para a certificação de alguns táxons inseridos em
complexos de espécies, como, por exemplo, os complexos P. albostipitatus e P. glaucotinctus
da seção Pluteus (capítulo II) e os complexos P. aureovanatus e P. riograndensis da seção
Celuloderma (capítulo V). O posicionamento da sequência de P. aff. cervinus (APÊNDICE
C) dentro do clado ‘aff. cervinus’, reconhecido por Justo et al. (2011b), pode revelar a
ocorrência de mais um táxon da seção Pluteus para o Brasil, porém, estudos multigênicos
também são indicados para o reconhecimento das espécies envolvidas nesse complexo.
Este trabalho não esgota o registro das espécies de Pluteus para o Brasil, ainda mais
considerando que as espécies reconhecidas estão restritas a 11 dos 26 estados brasileiros, mas
contribui significativamente para atualizar a conhecimento do gênero no país e para dar
subsídios para a condução de novos trabalhos de taxonomia e filogenia a partir de novas
coletas.
APÊNDICES
Apêndices - 224
Apêndice A - 225
APÊNDICE A – Lista comentada das espécies de Pluteus citadas para o Brasil
Táxons
Distribuição
P. aglaeotheles Berk. & Broome
AM
Referências bibliográficas e considerações
taxonômicas
Singer & Aguiar (1986)L como “P.
agloeotheles”
P. albostipitatus (Dennis) Singer
PR, RS, SP
Stijve & de Meijer (1993)L como P. cf.
albostipitatus, Pegler (1997)DI, Meijer
(2001, 2006)L, Wartchow et al. (2006)DC,
Menolli et al. (2010)DC, Menolli & Capelari
(2010)DC como P. densifibrillosus de acordo
com Justo et al. (2011b)
AM
Singer (1989)DC
P. aquosus Singer
RS
Wartchow et al. (2006)DC
P. atricapillus var. brasiliensis (Bres.)
RS
Raithelhuber (1991)L
SP
Menolli et al. (2010)DC, Justo et al. (2011a)L,
P. amazonicus Singer
B
Raithelh. [nom. inval.]
P. aureovenatus Menolli & CapelariB
Justo et al. (2011b)L
P. beniensis Singer
PR, RS
Stijve & de Meijer (1993)L como P. cf.
beniensis, Meijer (2006)L como P. cf.
beniensis, Wartchow et al. (2006)DC
P. brunneopictus Berk. & BroomeW
RS
Rick (1938, 1961)DI como “P. brunneopictus”, mas considerado P. umbrinoalbidus
ou P. rimosoaffinis por Singer (1959)
P. burserae Singer
RO
Singer (1959)DC
P. cervinus (Schaeff.) P. Kumm.W
RS
Rick (1938, 1961)DI, mas considerado P.
cervinus var. brasiliensis por Singer &
Digilio (1952) e posteriormente P.
xylophilus por Singer (1959)
P. cervinus var. brasiliensis Bres.B
RS
Bresadola (1920)DI, Singer & Digilio
(1952)DC, Singer (1959)DC como sinônimo
de P. xylophilus
B, W
SP
Hennings (1904b)DI, Pegler (1997)DI
P. cervinus var. patricius (Schaeff.) Quél. w
RS
Rick (1907)DI
P. chrysophlebius subsp. bruchii (Speg.)
PR
Stijve & de Meijer (1993)L como P.
P. cervinus var. griseoviridis Henn.
Singer
chrysophlebius subsp. bruchii var. bruchii,
Meijer (2006)L as P. chrysophlebius subsp.
bruchii var. bruchii, mas ambos
considerados por Menolli et al. (2010) como
prováveis P. sublaevigatus
Táxons
Distribuição
P. cristatulus RickB, W
RS
Apêndice A - 226
taxonômicas
Rick (1938, 1961)DI, mas considerado nomem
dubium por Singer (1959)
P. cubensis (Murrill) Dennis
MG, PR, SP
Stijve & de Meijer (1993)L, Pegler (1997)DI,
Meijer (2006)L, Rosa et al. (2003)L,
Xavier-Santos et al. (2004)L como P. cf.
cubensis, Rosa & Capelari (2009)L
P. densifibrillosus Menolli & CapelariB
SP
Menolli & Capelari (2010)DC, Justo et al.
(2011a)L, Justo et al. (2011b)L como
sinônimo de P. albostipitatus
P. diptychocystis Singer
SP
Pegler (1997)DI
P. dominicanus var. hyalinus Menolli &
SP
Menolli et al. (2010)DC
P. eucryphiae Singer
PR
Meijer (2006)L como P. cf. eucryphiae
P. eugraptus (Berk. & Broome) Sacc.
PR
Meijer (2006)L
P. exiguus (Pat.) Sacc.W
RS
Rick (1938, 1961)DI
P. exiguus var. venosus Rick [nom. inval.]B, W
RS
Rick (1961)L como “P. exiguus var. venosa”,
CapelariB
mas considerado por Singer (1954) como P.
phlebophorus sensu Lange e posteriormente
(Singer 1959) como provável P. jamaicensis
ou P. fluminensis
P. eximius (W. Saunders & W.G. Sm.) Sacc.
P. fallax Singer
W
RS
Rick (1919L, 1938DI, 1961DI)
RS
Singer (1954DC, 1956L) como P.
umbrinoalbidus somente para a coleção
Singer B110 (LIL), Singer (1959) DC
P. fibrillosus RickB,W
RS
Rick (1938, 1961)DI, mas considerado por
Singer (1959) como nomem dubium ou um
provável P. subfibrillosus
P. fibulatus Singer
PR, RS, SP
Singer & Digilio (1952)DC, Singer (1954)L,
Singer (1959)DC, Rick (1961)DI como P.
velatus de acordo com Singer (1954, 1959),
Raithelhuber (1991)L, Stijve & de Meijer
(1993)L, Pegler (1997)DI, Meijer (2006)L
P. fluminensis SingerB
PR, RJ, SP
Singer (1956)DC como P. variipes somente para
a coleção Singer B432 (F), Singer (1959)DC,
Stijve & de Meijer (1993)L, Meijer (2006)L
como P. cf. fluminensis, Pegler (1997)DI,
P. fuligineovenosus Horak
SP
P. glaucotinctus Horak
SP
Justo et al. (2011a, b)L
Táxons
P. glaucus Singer
Distribuição
PR
Apêndice A - 227
taxonômicas
Stijve & de Meijer (1993)DC como “Pluteus
sp. (aff. glaucus ” de acordo com Mei er
(2006), Stijve & de Meijer (1993)DC como P.
glaucus mas reidentificado como P.
salicinus por Meijer (2006), Stijve (1995)L,
Meijer (2006)L
P. globiger Singer
RS
P. granulatus Bres.W
RS
Rick (1930, 1938, 1961)DI, mas considerado
por Singer (1959) como um identificação
errada
P. harrisii Murrill
MG, PR, SP
Pegler (1997)DI, Meijer (2006)L como P. aff.
harrisii, Rosa & Capelari (2009)L, Menolli
& Capelari (2010)DC como P. puttemansii de
acordo com Justo et al. (2011b), Menolli et
al. (2010)DC
P. haywardii Singer
MG, PR, SP
Pegler (1997)DI, Meijer (2006)L como P. cf.
haywardii, Rosa & Capelari (2009)L
P. hispidulus (Fr.) GilletW
RS
Rick (1919L, 1938DI, 1961DI), mas considerado
por Singer (1959) como alguma outra
espécie da estirpe Fuliginosus
P. hylaeicola Singer
B
P. iguazuensis Singer
AM, PA
Singer (1989)DC
PR, RS
Meijer (2006)L como P. cf. iguazuensis,
Drechsler-Santos et al. (2007)L
P. jamaicensis Murrill
SP
P. laetifrons var. laetifrons (Berk. & M.A.
PR
Meijer (2006)L, Meijer (2009)L
P. laetifrons var. floridae Singer
PR
Meijer (2006)L
P. leoninus (Schaeff.) P. KummW
RS
Curtis) Sacc.
Singer (1959) como um provável P.
chrysophlebius ssp. bruchii ou uma outra
espécie de basidioma amarelo
B, W
P. leptonia Rick
RS
Rick (1938, 1961)DI como “Pluteus
Leptonia”, mas considerado por Singer
(1959) como uma provável Nolanea howellii
Peck
P. longistriatus (Peck) Peck
SP
Menolli & Capelari (2010)DC, Justo et al.
(2011a, b)L, Gugliotta et al. (2012) L
Táxons
Distribuição
P. melanodon Sacc.
RS
Apêndice A - 228
taxonômicas
Singer (1959) como alguma outra espécie de
Pluteus com borda da lamella pigmentada ou
ainda alguma espécie de Acurtis
(Rhodophyllus)
P. melanopotamicus Singer
B
AM
Singer & Aguiar (1986)L nom. nud., Singer
(1989)DC
P. minutes Pat.
BA
Singer (1989)L
P. nanus (Pers.) P. Kumm.W
RS
por Singer (1959) como uma identificação
duvidosa
P. nanus var. podospileus (Sacc. & Cub.)
RS
RickW
Rick (1938)DI, Rick (1961)DI como “P.
nanus var. podospilea”
P. neophlebophorus Singer
PR
Meijer (2006)L como P. aff. neophlebophorus
P. nigrolineatus Murrill
RS
P. paraensis Singer B
PA
Singer (1973)DC
P. pellitus (Pers.) P. Kumm.W
RS
P. viscidulus por Singer (1954, 1959)
P. phlebophorus (Ditmar) P. Kumm.
W
RS
Rick (1938, 1961)DI, mas considerado por
Singer (1959) um provável P. jamaicensis
P. pluvialis Singer
SP
Pegler (1997)DI
P. polycystis Singer
PR
Meijer (2006)L as P. cf. polycystis
P. pulverulentus var. pseudonanus Singer
PR
Stijve & de Meijer (1993)L, Meijer (2006)L
P. puttemansii Menolli & CapelariB
SP
Menolli & Capelari (2010)DC, Meijer (2010)L,
Justo et al. (2011a)L, Justo et al. (2011b)L
como sinônimo de P. harrisii
P. riberaltensis var. conquistensis Singer
SP
Menolli et al. (2010)DC, Justo et al. (2011a)L,
Justo et al. (2011b)L
P. rimosellus Singer
RS
Singer (1954)L, mas reidentificado como P.
subfibrillosus em Singer (1956, 1959)
P. rimosoaffinis Singer
PR, SP
Pegler (1997)DI, Meijer (2006)L como P. cf.
rimosoaffinis
P. riograndensis Singer
B
RS
Singer (1954DC, 1956L, 1959DC), Raithelhuber
(1991)L
P. salicinus (Pers.) P. Kumm.
PR
Stijve & de Meijer (1993)DC como P. glaucus
according to Meijer (2006), Meijer (2006)L
P. scruposus Henn.B, W
MT
Hennings (1900)DI, mas considerado por Singer
(1959) como sinônimo de Oudemansiella
canarii (Jungh.) Höhn.
Táxons
Distribuição
Apêndice A - 229
taxonômicas
P. sensitivus RickB, W
RS
Rick (1930, 1938, 1961)DI
P. sergii Singer
SP
Pegler (1997)DI
P. straminellus RickB, W
RS
Rick (1961)DI
P. subfibrillosus SingerB
PR, RJ, RS
Singer (1954)L como P. rimosellus de acordo
com (Singer 1956, 1959), Singer (1956,
1959)DC, Meijer (2009)L
P. sublaevigatus (Singer) Menolli & Capelari
SP
P. termitum Henn.B, W
AM
Hennings (1904a)DI como “Pluteus ?
termitarum”, mas considerado por Saccardo
& Trotter (1912) como sinônimo de Collybia
eurhiza (Berk.) Höhn. ou uma provável
Lepiota sp. por Singer (1959)
P. thomsonii (Berk. & Br.) Dennis
PR, RS
Wartchow et al. (2004)DC, mas considerado por
Rodríguez et al. (2009) como um provável
P. neotropicalis Rodr.-Alcánt., Meijer
(2006)L como P. aff. thomsonii
P. umbrinoalbidus SingerB
MG, PR, RS, Singer (1954DC, 1956L, 1959DC), Raithelhuber
SP
(1991)L, Meijer (2006, 2009)L como “P.
cf. umbrinoalbus”, osa & Capelari
(2009)L, Menolli & Capelari (2010)DC
P. umbrosus (Pers.) P. Kumm.W
RS
Rick (1919L, 1938DI, 1961DI)
P. variipes Singer
RJ
Singer (1956)DC, mas reidentificado como P.
fluminensis em Singer (1959)
Singer (1984)L
P. varzeae nom. nud.
AM (?)
P. varzeicola SingerB
AM
Singer (1984)L nom. nud., Singer (1989)L
P. velatus RickB, W
RS
Rick (1961)DI, Singer (1954L, 1959DC) como P.
fibulatus
P. viscidulus Singer
RS
Rick (1930, 1938, 1961)DI como P. pellitus
de acordo com Singer (1954, 1959), Singer
(1954L, 1959DC)
P. xylophilus (Speg.) Singer
PR, RS, SP
Bresadola (1920)DI como P. cervinus var.
brasiliensis, Rick (1938, 1961)DI como P.
cervinus, Singer & Digilio (1952)DC como
P. cervinus var. brasiliensis, Singer
(1959)DC, Grandi et al. (1984)L, Stijve & de
Meijer (1993)L, Pegler (1997)DI,
Sobestiansky (2005)L como P. cf.
xylophilus, Meijer (2006L, 2009DC), Menolli
et al. (2010)DC, Justo et al. (2011ab)L
Táxons
Distribuição
Apêndice A - 230
taxonômicas
P. xylophilus var. tucumanensis Singer
PR
Stijve & de Meijer (1993)L
Pluteus sp.
MG
Rosa et al. (2003)L, Rosa & Capelari (2009)L
Pluteus sp.
RS
Pereira (1984)L
Pluteus sp.
RO
Capelari & Maziero (1988)L
Pluteus sp.
RS
Guerrero & Homrich (1999)L
Pluteus sp.
SP
Menolli et al. (2010)L
 = referência que não indicou a coleção estudada e/ou o herbário no qual o material foi depositado. B = espécie com coleção tipo coletada no
Brasil; L = referência que mencionou a espécie apenas em lista e/ou sem descrição morfológica; DC = referência que publicou descrição
completa para espécie; DI = referência que publicou descrição morfológica incompleta.
Apêndice B - 231
APÊNDICE B – Chave de identificação para as espécies de Pluteus
ocorrentes no Brasil*†
1a.
Pleurocistídios metuloides, a maioria com prolongamentos apicais, raramente de parede
levemente espessada ou não metuloide; superfície pilear indiferenciada composta por
hifas prostradas; ansas presentes ou ausentes ..................................... (Seção Pluteus) 2
1b.
Pleurocistídios de parede fina, raramente de parede levemente espessada ou quando
metuloides sem prolongamentos apicais; superfície pilear epitelial, himenidérmica,
tricodérmica ou indiferenciada composta por hifas prostradas; ansas ausentes ................
........................................................................ (Seções Celluloderma e Hispidoderma) 8
2a.
Ansas presentes ............................................................................................................... 3
2b.
Ansas ausentes ................................................................................................................ 5
3a.
Borda da lamela distintamente pigmentada; queilocistídios pigmentados .. P. hylaeicola
3b.
Borda da lamela concolor com a face; queilocistídios sem pigmentação ....................... 4
4a.
Pleurocistídios metuloides, de parede espessada e ápice geralmente com dois a quatro
prolongamentos apicais, raramente sem prolongamentos e com ápice agudo ou
arredondado; queilocistídios exclusivamente clavados com longo pedicelo basal;
basidiósporos globosos a levemente elípticos (raramente elípticos) ............. P. fibulatus
4b.
Pleurocistídios de parede fina ou ocasionalmente levemente espessada no ápice, ápice
arredondado ou truncado e sem prolongamentos apicais; queilocistídios usualmente
mucronados ou fusiformes, basidiósporos subglobosos a elípticos (raramente
cilíndricos) .............................................................................................. P. glaucotinctus
5a.
Píleo inteiramente branco ........................... P. petasatus (forma albina = P. viscidulus)
5b.
Píleo pigmentado em tons de marrom ............................................................................ 6
6a.
Pleurocistídios metuloides com ápice de três tipos: com dois a seis prolongamentos bem
desenvolvidos, agudo e sem prolongamentos ou sem prolongamentos e com ápice
trucado. Basidiósporos < 5.0 µm de largura ................................................ P. xylophilus
6b.
Pleurocistídios metuloides ou de parede levemente espessada (raramente de parede
fina), com ou sem prolongamentos apicais, mas nunca com ápice agudo. Basidiósporos
> 5.0 µm de largura ......................................................................................................... 7
7a.
Basidiósporos levemente elípticos a elípticos (raramente subglobosos); pleurocistídios
distintamente metuloides, com dois a cinco prolongamentos apicais bem desenvolvidos,
ocasionalmente com dois a quatro prolongamentos curtos e pouco desenvolvidos, ou
ainda sem prolongamentos e com ápice truncado a arredondado ..................... P. harrisii
* Táxons que precisam de novas coletas para certificar sua ocorrência no Brasil
7b.
Apêndice B - 232
Basidiósporos globosos a levemente elípticos (raramente elípticos ou cilíndricos);
pleurocistídios de parede fina a levemente espessada, usualmente sem prolongamentos
apicais e com ápice truncado a arredondado, raramente como metuloides (parede ≤ 1.0
µm de espessura) e com dois a quatro prolongamentos apicais ............. P. albostipitatus
8a.
Superfície pilear tricodérmica ou himenidérmica de elementos longos (Qm > 3). ...........
.................................................................................................... (Seção Hispidoderma) 9
8b.
Superfície pilear indiferenciada composta por hifas prostradas ou epitelial /
himenidérmica de elementos curtos (Qm < 3), com ou sem elementos cistidioides
entremeados ............................................................................. (Seção Celluloderma) 17
9a.
Píleo branco a branco-rosado ........................................................................ P. aquosus*
9b.
Píleo distintamente pigmentado: cinza, marrom ou enegrecido ................................... 10
10a. Pleurocistídios ausentes ............................................................................ P. maculosipes
10b. Pleurocistídios presentes ............................................................................................... 11
11a. Pleurocistídios de parede espessada e pigmentados; queilocistídios pigmentados e
dimórficos: I) de parede fina e com ápice arredondado, clavados, ventricosos ou
utriformes; II) de parede espessada e fusiforme com ápice agudo ........ P. neochrysaegis
11b. Pleuro- e queilocistídios diferentes dos descritos acima .............................................. 12
12a. Borda da lamela distintamente pigmentada ................................................ P. chusqueae
12b. Borda da lamela concolor com a face ........................................................................... 13
13a. Pleurocistídios totalmente hialinos e sem conteúdo ............................... P. argentinensis
13b. Pleurocistídios com conteúdo interno incolor a levemente amarelado ......................... 14
14a. Margem do píleo não sulcada ........................................................................ P. velutinus
14b. Margem do píleo distintamente sulcada ....................................................................... 15
15a. Píleo com centro levemente venoso-rugoso ................................................ P. varzeicola
15b. Píleo com centro fibriloso a punctado-esquamuloso .................................................... 16
16a. Píleo profundamente sulcado, ao menos até a metade do píleo, e fibriloso-granuloso
apenas no centro, pleurocistídios até 114 µm de comprimento ............... P. longistriatus
16b. Píleo distintamente fibriloso-rimoso, pleurocistídios até 65 µm de comprimento ............
...................................................................................................................... P. rimosellus
17a. Superfície pilear indiferenciada composta por hifas prostradas; pleurocistídios
exclusivamente de parede fina ...................................................................................... 18
17b. Superfície pilear epitelial ou himenidérmica, com ou sem elementos cistidioides
entremeados; pleurocistídios de parede fina ou levemente espessada na porção apical 24
18a. Pleurocistídios raros ou ausentes .................................................................................. 19
18b. Pleurocistídios comumente presentes ........................................................................... 20
Apêndice B - 233
19a. Margem do píleo estriada-sulcada; pleurocistídios ausentes; queilocisítios raros
(fusoide-ventricosos a levemente lageniformes) .................................... P. karstedtianus
19b. Margem do píleo não estriada ou sulcada; pleurocistídios raros (ventricosos a
lageniformes ou alongado-clavados); queilocistídios frequentes e de formas muito
variadas (clavado, ampuláceo, fusiforme, capitado ou esferopedunculado) .....................
........................................................................................................... P. hispidulussimilis
20a. Pleurocistídios sem qualquer ornamentação externa .................................................... 21
20b. Pleurocistídios com incrustação resinosa apical ou com colar médio-apical fortemente
aderido à parede ............................................................................................................ 22
21a. Píleo levemente rimoso, margem não sulcata ou estriada; estipe esbranquiçado mas
coberto com pequenas fibrilas marrom-acinzentadas ........................................................
................................................................................... P. riberaltensis var. conquistensis
21b. Píleo radialmente fibriloso, margem levemente estriada; estipe completamente branco ..
......................................................................................................................... P. cubensis
22a. Basidiósporos globosos a subglobosos; pleurocistídios com incrustação resinosa
amarelada no ápice formando uma linha esférica bem delimitada ............. P. aureolatus
22b. Basidiósporos subglobosos a elípticos; pleurocistídios sem incrustação resinosa
amarelada no ápice mas com colar médio-apical fortemente aderido à parede ............ 23
23a. Pleurocistídios exclusivamente sem conteúdo e com colar liso aderido à parede .............
................................................................................................................ P. diptychocystis
23b. Pleurocistídios com conteúdo incolor ou levemente pigmentado e com colar estriado
aderido à parede ......................................................................................... P. striatocystis
24a. Superfície
pilear
com
elementos
cistidioides
entremeados;
pleurocistídios
exclusivamente de parede fina ...................................................................................... 25
24b. Superfície pilear sem elementos cistidioides entremeados; pleurocistídios de parede fina
ou levemente espessada na porção apical ..................................................................... 33
25a. Pleurocistídios ausentes (ou muito raros) ..................................................................... 26
25b. Pleurocistídios sempre presentes .................................................................................. 29
26a. Borda da lamela distintamente pigmentada; queilocistídios fortemente pigmentados ......
............................................................................................................ P. brunneocrinitus
26b. Borda da lamela concolor com face; queilocistídios sem pigmentação ....................... 27
27a. Píleo distintamente reticulado-venoso; elementos da superfície pilear, queilo- e
pleurocistídios (quando presentes) frequentemente com uma longa projeção apical (até
62 µm de comprimento) ............................................................................... P. thomsonii
Apêndice B - 234
27b. Píleo distintamente fibriloso; elementos da superfície superior e queilocistídios sem
qualquer projeção apical; pleurocistídios completamente ausentes ............................. 28
28a. Queilocisítios levemente ventricosos, subglobosos ou esferopedunculados [25–34(–39)
× 12.5–21(–26) µm]; caulocistídios clavados a fusoide-ventricosos (37–75 × 8.7–21
µm) ................................................................................................................... P. crinitus
28b. Queilocistídios clavados a ventrico-clavados ou levemente lageniformes (37–75 × 12.5–
30 µm); caulocistídios aparentemente ausentes mas contexto do estipe constituído de
elementos terminais diferenciados, de formato irregular, de parede levemente espessada
a espessada e às vezes com pontos de incrustação (28–50 × 3.7–8.7 µm) ........................
................................................................................................................... P. necopinatus
29a. Queilocistídios muito maiores em comprimento que os pleurocistídios (até 3,5 ×),
ambos fortemente pigmentados ....................................................... P. anomocystidiatus
29b. Queilocisídios de tamanho similar aos pleurocistídios; pleurocistídios sem nenhum
pigmentação; queilocistídios pigmentados ou não ....................................................... 30
30a. Margem do píleo fortemente sulcada (não meramente estriada por transparência) ..... 31
30b. Margem do píleo não sulcada ou estriada (ou levemente estriada por transparência) . 32
31a. Píleo diminuto (até 3 mm diam.); queilocistídios completamente sem conteúdo;
elementos cistidioides da superfície pilear geralmente com ápices arredondados ............
................................................................................................................... P. amazonicus
31b. Píleo maior que 3 mm diam.; queilocistídios às vezes com conteúdo marrom pálido;
elementos cistidioides da superfície pilear normalmente com ápices subagudos .............
........................................................................................................................ P. burserae
32a. Píleo fortemente rugoso-venoso em toda a extensão ou somente no centro;
queilocistídios às vezes com conteúdo marrom pálido .................................... P. eludens
32b. Píleo liso, às vezes rimoso mas nunca rugoso-venoso; queilocistídios completamente
sem conteúdo ...................................................................................... P. umbrinoalbidus
33a. Píleo, lamelas ou estipe com tonalidades de cor amarela ou alaranjada ....................... 34
33b. Píleo, lamelas ou estipe sem qualquer tonalidade de cor amarela ou alaranjada,
normalmente em tons de marrom ou bege .................................................................... 37
34a. Pleuro- e queilocistídios frequentemente de parede espessada (até 1.2 µm) e com uma
capa apical bem definida e fortemente aderida à parede .................................. P. crassus
34b. Pleuro- e queilocistídios exclusivamente de parede fina, sem capa apical aderida à
parede ou às vezes apenas com mucilagem amorfa no ápice dos pleurocistídios ........ 35
35a. Píleo nunca puramente amarelo ou alaranjado, mas sim amarelado com tons de marrom;
células da superfície pilear em sua maioria pigmentada com conteúdo marrom claro .....
Apêndice B - 235
......................................................................................................................... P. globiger
35b. Píleo completamente amarelo ou com tons alaranjados; células da superfície pilear
majoritariamente sem nenhuma pigmentação ............................................................... 36
36a. Píleo completamente amarelo, sem tons alaranjados ............................. P. sublaevigatus
36b. Píleo fortemente amarelado, coberto por veias alaranjadas em toda sua extensão ............
................................................................................................................. P. aureovenatus
37a. Pleurocistídios ausentes (ou muito raros) ..................................................................... 38
37b. Pleurocistídios presentes ............................................................................................... 39
38a. Margem do píleo branca e fortemente plissada-sulcada; basídios dimórficos ..................
..................................................................................................................... P. cebolinhae
38b. Margem do píleo concolor ou levemente mais pálida que o centro, sulcada-estriada mas
não distintamente plissada; basídios uniformes ................................. P. paucicystidiatus
39a. Píleo bege ou marrom claro; células da superfície pilear sem pigmentação .....................
........................................................................................... P. dominicanus var. hyalinus
39b. Píleo distintamente marrom; células da superfície pilear sempre pigmentadas ........... 40
40a. Borda da lamela distintamente pigmentada; queilocistídios sempre pigmentados ....... 41
40b. Borda da lamela concolor com a face ou às vezes irregularmente ou fracamente
pigmentada; queilocistídios sem conteúdo ou fracamente pigmentados ...................... 42
41a. Pleurocistídios pigmentados ................................................................ P. fusconigricans
41b. Pleurocistídios sem pigmentação .................................................................... P. homolae
42a. Pleuro- e/ou queilocistídios fracamente pigmentados .................................................. 43
42b. Pleuro- e queilocistídios sempre sem pigmentação ...................................................... 44
43a. Píleo distintamente rimoso e craquelado; margem da lamela concolor com a face ..........
................................................................................................................. P. rimosoaffinis
43b. Píleo inteiro, não rimoso ou craquelado; margem da lamela às vezes irregularmente ou
fracamente pigmentada .......................................................................... P. riograndensis
44a. Centro do píleo liso, mesmo quando seco .................................................................... 45
44b. Centro do píleo distintamente rugoso-venoso ou pelo menos levemente rugoso quando
seco ............................................................................................................................... 46
45a. Píleo radialmente rimoso; estipe completamente branco ................................. P. fallax*
45b. Píleo distintamente sulcado-estriado, mas não rimoso; estipe creme-esbranquiçado com
a base levemente marrom .......................................................................... P. iguazuensis
46a. Basidiósporos na maioria geometricamente globosos ou subglobosos ......... P. sapiicola
46b. Basidiósporos na maioria subglobosos a levemente elípticos, nunca perfeitamente
globosos ........................................................................................................................ 47
Apêndice B - 236
47a. Centro do píleo distintamente rugoso-venoso, margem do píleo não sulcada ...................
.................................................................................................................... P. fluminensis
47b. Centro do levemente rugoso apenas quando seco, margem do píleo distintamente
sulcada ........................................................................................................ P. paraensis*
Apêndice C - 237
APÊNDICE C – Lista das coleções brasileiras de Pluteus depositadas em herbário
Considerações
Táxons*‡
taxonômicas/ dados
Coleção / Origem
moleculares
Referências
bibliográficas
P. aglaeotheles
Não localizado
Brasil, AM, Tarumãzinho (F?)
Singer & Aguiar
(1986)
P. albostipitatus
Identificação
confirmada
A. Batista s.n., 19/III/1980, Brasil, RS,
São Francisco de Paula, Fl. Nac. do
IBDF (ICN6834)
Pereira (1984)
como Pluteus sp.
P. albostipitatus
Identificação
confirmada
A.A.R. de Meijer MANa-2629,
20/IV/1993, Brasil, PR, Mandirituba
(MBM371810)
Meijer (2006)
P. albostipitatus
Identificação
confirmada
A.A.R. de Meijer PAc-2744, 27/V/1993, Meijer (2006)
Brasil, PR, Paranaguá, Mundo Novo
(MBM371819)
P. albostipitatus
Identificação
confirmada
A.A.R. de Meijer ANc-4219,
24/V/2003, Brasil, PR, Antonina,
Reserva Natural do Rio Cachoeira
(MBM371812)
P. albostipitatus
Identificação
confirmada na
duplicata
D.N. Pegler 3890, 30/I/1987, Brasil, SP, Pegler (1997)
Mogi-Guaçu [SP214374 = K(M)41612 como P. sergii
como P. sergii]
P. albostipitatus
Identificação
confirmada
Mogi-Guaçu [K(M)41612 = SP214374 como P. sergii
como P. sergii]
P. albostipitatus
Insuficiente para
estudo
Campos do Jordão [SP214367 =
K(M)41606]
P. albostipitatus
Identificação
confirmada (Seq. ITS:
FJ816661, Seq. LSU:
FJ816647)
F. Karstedt & M. Capelari FK782,
23/X/2006, Brasil, SP, São Paulo,
Parque Estadual da Cantareira, Núcleo
Engordador (SP393714)
Menolli et al.
(2010)
P. albostipitatus
Identificação
confirmada
F. Karstedt et al. FK1061, 31/I/2008,
Brasil, SP, São Paulo, Parque Estadual
da Cantareira, Núcleo Engordador
(SP417353)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
R. Singer B11231, 12/VI/1978, Brasil,
BA, Itabuna (INPA82449)
Dados não
publicados
P. albostipitatus
Identificação
JQ065033)
F. Karstedt FK1664, 07/XI/2010, Brasil, Dados não
PR, Foz do Iguaçu, Parque Nacional do publicados
Iguaçu, Trilha do Hidrômetro
(SP416666)
P. albostipitatus
Identificação
JQ801373)
F. Karstedt FK1891, 20/I/2011, Brasil,
PA, Belém, Ilha do Combu (SP416744)
*
Táxons em negrito representam coleções examinadas neste trabalho.
Meijer (2006) as P.
cf. harrisii
Dados não
publicados
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 238
Referências
bibliográficas
P. albostipitatus
Identificação
JQ065032)
F. Karstedt FK1973, 18/III/2011, Brasil, Dados não
MG, Alto Caparaó, Parque Nacional do publicados
Caparaó (SP416665)
P. albostipitatus
Identificação
confirmada
F. Karstedt FK1598, 30/VII/2010,
Brasil, SC, Gaspar, Reserva Particular
do Patrimônio Natural Figueira Branca
(SP417351)
Dados não
publicados
P. albostipitatus
Identificação
JQ801375)
F. Karstedt & R. Karstedt FK1799,
25/I/2011, Brasil, SC, Blumenau,
Parque Nacional Serra do Itajaí, Parque
das Nascentes, Trilha da 3ª Vargem
(SP417346)
Dados não
publicados
P. albostipitatus.
Identificação
confirmada
25/I/2011, Brasil, SC, Blumenau,
Parque Nacional Serra do Itajaí, Parque
das Nascentes, Trilha da 3ª Vargem
(SP417352)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
F. Karstedt & J.J.S. Oliveira. FK1909,
01/III/2011, Brasil, SP, São Paulo,
Parque Estadual das Fontes do Ipiranga
(SP417125)
Dados não
publicados
P. albostipitatus
Identificação
confirmada (Seq. ITS
disponível para
depósito)
F. Karstedt FK2147, 06/I/2012, Brasil,
SC, Gaspar, RPPn Figueira Branca
(SP417720)
Dados não
publicados
P. albostipitatus
Insuficiente para
estudo
F. Wartchow & J.C. Budke 039,
22/V/2002, Brasil, RS, Santa Maria,
Morro do Elefante (SMDB9712)
Wartchow et al.
(2006)
P. albostipitatus
Identificação
confirmada
G. Guzman et al. 22947., 17/XI/1982,
Brasil, SP, Cananeia, Ilha do Cardoso,
Restinga (SP178032 como P. cubensis)
Pegler (1997)
como P. cubensis
P. albostipitatus
Identificação
confirmada
J. Medeiros s.n., 03/X/1984, Brasil, RS,
Guaíba, Horto Florestal da CEEE
(ICN102101)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
J. Rick s.n., 1944, Brasil, RS, São
Salvador (PACA20770 - holótipo de P.
sensitivus)
Rick (1961) como
P. sensitivus
P. albostipitatus
Identificação
confirmada
J. Rick s.n., Brasil (BPI770882 como P.
murinus)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
M.S. Rother & B.M.A. Severo,
30/X/2003, Brasil, RS, Passo Fundo,
UPF (RSPF284)
Dados não
publicados
P. albostipitatus
Identificação
FJ816656)
Menolli Jr. et al. NMJ128, 24/VII/2007, Menolli et al.
(2010)
(SP393713)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 239
Referências
bibliográficas
P. albostipitatus
Identificação
HM562159)
Menolli Jr. et al. NMJ129, 24/IV/2007,
(SP393696 - holótipo de P.
densifibrillosus)
Menolli &
Capelari (2010),
Justo et al. (2011
a, b) como P.
densifibrillosus
P. albostipitatus
Identificação
confirmada
R. Maziero s.n., 22/I/2007, Brasil, MG,
Nova Ponte, Fazenda Caxuanã
(SP417354)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
(SP417355)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
(SP417356)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
(SP417357)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
(SP417358)
Dados não
publicados
P. albostipitatus
Identificação
confirmada
(SP417359)
Dados não
publicados
P. albostipitatus
Identificação
JQ065031)
R. Singer B12426, 31/III/1983, Brasil,
AM, Igarapé do Tarumãzinho (Fholótipo de P. melanopotamicus)
Singer (1989)
como P.
melanopotamicus
P. albostipitatus
Insuficiente para
estudo
Pegler 3868, 28/I/1987, Brasil, SP,
Campos do Jordão K(M)41606
(=SP214367)
Pegler (1997)
P. albostipitatus
Não localizado
A.A.R. de Meijer CUc-1557, Brasil, PR, Meijer (2006)
Curitiba, Capão da Imbuia quarter,
Museu de História Natural Capão da
Imbuia, Forest (MBM)
P. albostipitatus
Não localizado
A.A.R. de Meijer REb, 14/V/1992,
Brasil, PR, Reserva do Iguaçu, Segredo
(MBM)
Meijer (2006)
P. amazonicus
Não localizado
R. Singer B 10187, 10/X/1977, Brasil,
AM, Estrada Manaus-Itacoatiara, 30
Km N de Manaus, Embrapa
(INPA77269 - holótipo)
Singer (1989)
P. cf. angustiporus
Identificação incerta
A.A.R. de Meijer SJf-4113, 31/X/2002,
Brasil, PR, São José dos Pinhais,
Contenda (MBM371827)
Meijer (2006)
como P. xylophilus
P. anomocystidiatus sp.
nov.
Identificação
confirmada
A.A.R. de Meijer DN-3378, 18/III/1996, Dados não
Brasil, PR, Diamante do Norte,
publicados
Ecológica do Caiuá (MBM371820 holótipo)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 240
Referências
bibliográficas
P. anomocystidiatus sp.
nov.
Identificação
confirmada
07/III/2009, Brasil, PR, Antonina,
(MBM371814)
Dados não
publicados
P. aquosus
Insuficiente para
estudo
V.G. Cortez 053/01, 24/XII/2001,
Brasil, RS, Santa Maria, Morro do
Elefante (SMDB9263)
Wartchow et al.
(2006)
P. argentinensis
Identificação
confirmada
L.H. Rosa & R.O. Morais PERD76,
10/I/2000, Brasil, MG, Marliéria,
Parque Estadual do Rio Doce
(SP307730 = BHCB68363 as P.
cubensis)
Rosa & Capelari
(2009) como P.
cubensis
P. argentinensis
Identificação
disponível para
depósito)
F. Karstedt et al. FK1903, 01/III/2011,
das Fontes do Ipiranga (SP417462)
Dados não
publicados
P. argentinensis
Identificação
confirmada
F. Karstedt et at. FK1915, 03/III/2011,
Dados não
publicados
P. aureolatus sp. nov.
Identificação
disponível para
depósito)
F. Karstedt FK2084, 29/VI/2011, Brasil, Dados não
AM, Manaus, Campus Aleixo I
publicados
(INPA241273 - holótipo)
P. aureolatus sp. nov.
Identificação
disponível para
depósito)
N.K. Ishikawa NKI7, 09/II/2011, Brasil, Dados não
AM, Manaus, INPA, Campus Aleixo I
publicados
(INPA239965)
P. aureovenatus
Identificação
FJ816663, Seq. LSU:
FJ816649)
F. Karstedt & L.A.S. Ramos FK1045,
Menolli et al.
24/I/2008, Brasil, SP, São Paulo, Parque (2010)
Estadual das Fontes do Ipiranga
(SP393697 - holótipo)
P. aureovantus
Identificação
disponível para
depósito)
Brasil, RJ, Parque Nacional Serra dos
Órgãos, Sede Guapimirim, Poço da
Ponte Velha (SP416735)
Dados não
publicados
P. avellaneus
Indisponível para
estudo
SMDB9193
Dados não
publicados
P. cf. beniensis
Não localizado
A.A.R. de Meijer CUb, 31/XII/1991,
Brasil, PR, Curitiba, Parque Barigui
(MBM)
Meijer (2006)
P. brunneocrinitus sp.
nov.
Identificação
disponível para
depósito)
M. Capelari & L.A.S. Ramos MC4535,
25/XI/2009, Brasil, SP, São Paulo,
Parque Estadual da Cantareira
(SP417458 – holótipo)
Dados não
publicados
P. brunneopictus
Provável P.
tucumanus
J. Rick s.n., 1936, Brasil, RS, Santa
Maria (PACA14522)
Rick (1961)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 241
Referências
bibliográficas
P. brunneopictus
Provável P. cubensis
Maria (PACA14531)
Rick (1961)
P. brunneopictus
Não localizado
Maria (PACA14532)
Dados não
publicados
P. burserae
Identificação
confirmada
R. Singer B1805, 10/III/1956, Brasil,
RO, Guajará Mirim (LIL)
Singer (1959)
P. cebolinhae sp. nov.
Identificação
disponível para
depósito)
C.A. Canavese & F. Karstedt CAC39,
20/VI/2007, Brasil, SP, Santo André,
Reserva Biológica de Paranapiacaba
(SP417455 – holótipo)
Dados não
publicados
P. cervinus
Provável P.
angustisporus
J. Rick s.n., 1920, Brasil, RS, Arroio do
Meio (BPI770858)
Dados não
publicados
P. cervinus
Provável P.
angustisporus
Meio (SP33926)
Dados não
publicados
P. cervinus
Indisponível para
estudo
Rick s.n., 1906, Brasil (FH00301661)
Dados não
publicados
P. cervinus
Indisponível para
estudo
Rick s.n., 1901, Brasil, RS
(FH00301662)
Dados não
publicados
P. cervinus
Indisponível para
estudo
Rick s.n., 1928, Brasil, SC
(FH00301663)
Dados não
publicados
P. cervinus
Indisponível para
estudo
Rick s.n., 1928, Brasil, SC
(FH00301664)
Dados não
publicados
P. cervinus
Indisponível para
estudo
Rick s.n., 1928, Brasil, RS
(FH00301665)
Dados não
publicados
P. aff. cervinus
Sequência ITS
disponível para
depósito (posicionada
no clado ‘aff.
cervinus’ de Justo et
al. 2011b)
M.S. Rother & B.M.A. Severo
16/I/2004, Brasil, RS, Passo Fundo,
Jaboticabal (RSPF323 como P.
spinulosus)
Dados não
publicados
P. cervinus var.
brasiliensis
Não localizado
J. Rick no. 412, Brasil, RS, São
Leopoldo (O ?)
Bresadola (1920)
P. cervinus var.
griseoviridis
Provável P.
angustisporus
A. Puttemans 871, III/1903, Brasil, SP,
São Paulo, Serra da Cantareira
(SP141797 - holótipo de P. cervinus
var. griseoviridis)
Hennings (1904b),
Pegler (1997)
P. chrysophlebius subsp.
bruchii
Não localizado
A.A.R. de Meijer PAa-1785, Brasil, PR, Meijer (2006)
Paranaguá, Ilha do Mel (MBM)
P. chusqueae stat. nov.
Identificação
confirmada
A.A.R. de Meijer & M.A.L. de A.
Meijer (2006)
Amazonas COa-3748, 10/IV/2000,
como P. eugraptus
Brasil, PR, Colombo, Embrapa Florestas
(MBM371805)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 242
Referências
bibliográficas
P. crassus sp. nov.
Identificação
confirmada
Amazonas COa-4057, 21/III/2001,
(MBM371825 - holótipo)
Meijer (2006)
como P.
chrysophlebius
subsp. bruchii
P. crassus sp. nov
Identificação
confirmada
(MBM371801)
Meijer (2006)
como P. cf.
iguazuensis
P. crinitus sp. nov.
Identificação
disponível para
depósito)
F. Karstedt FK2064, 27/VI/2011, Brasil, Dados não
AM, Manaus, INPA, Campus Aleixo I
publicados
(INPA241254 – holótipo; SP445855 isótipo)
P. cubensis
Identificação
confirmada
A.A.R. de Meijer MANa-2631,
20/IV/1993, Brasil, PR, Mandirituba
(MBM371809)
Meijer (2006)
P. cubensis
Identificação
HM562161)
C. Puccinelli et al. CP233, 23/III/2007,
Brasil, SP, Iporanga, PETAR, Núcleo
Ouro Grosso (SP394389)
Justo et al. (2011a,
b) como Pluteus
sp. V
P. cubensis
Identificação
confirmada
Mogi-Guaçu [K(M)41607 = SP214310]
P. cubensis
Não localizado (ident.
confirmada na
duplicata)
Mogi-Guaçu [SP214310 = K(M)41607]
P. cubensis
Entolomataceae –
Provável Rhodocybe
sp.
I.C. Oliveira 286, 23/VII/1990, Brasil,
PB, João Pessoa, Campus I da UFPB
(JPB17533 as P. cubensis)
Dados não
publicados
P. cubensis
Identificação
confirmada
M. Capelari et al. MC1367, 11/V/1987,
Brasil, RO, Jaru, Reserva Biológica do
Jaru, margem direita do rio Ji-Paraná
(SP212106)
Dados não
publicados
P. cubensis
Identificação
confirmada
Jaru, margem direita do rio Ji-Paraná,
próximo à ilha 7 de setembro
(SP214560 as P. cf. cubensis)
Dados não
publicados
P. cubensis
Insuficiente para
estudo
M. Capelari MC248, 29/I/1985, Brasil,
SP, Cananeia, Ilha do Cardoso, entre
morro dos Três Irmãos e Ipanema
(SP193836)
Pegler (1997)
P. cubensis
Não localizado
A.A.R. de Meijer MAg, 2/VI/1991,
Brasil, PR, Morretes, Rio
Nhundiaquara, eastern part of Caminho
do Itupava, train satations Marumbi,
Eng. Lange and Véu da Noiva (MBM)
Meijer (2006)
P. cf. cubensis
Indisponível para
estudo
S.X. Santos 320, Brasil, SP, São José do Xavier-Santos et
Rio Preto, Estação Ecológica do
al. (2004)
Noroeste Paulista (SJRP)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 243
Referências
bibliográficas
P. dominicanus var.
hyalinus
Identificação
FJ816665, Seq. LSU:
FJ816651)
(SP393695 - holótipo)
Menolli et al.
(2010)
P. dissimilis (nom. in
herb.)
Provável P.
angustisporus
leg. Steffen s.n., det. J. Rick, 1929,
Dados não
Brasil, RS, São Leopoldo (PACA14517) publicados
P. diptychocystis
Identificação
disponível para
depósito)
Menolli Jr. et al. NMJ184, 14/VII/2010, Dados não
Brasil, RN, Baía Formosa, Reserva
publicados
Particular do Patrimônio Natural Mata
Estrela (SP445829)
P. diptychocystis
Identificação
confirmada
SP, Cananeia, Parque Estadual da Ilha
do Cardoso, entre Morro Três Irmãos e
Ipanema (SP193821)
Dados não
publicados
P. egregius (nom. in
herb.)
Provável P.
angustisporus
J. Rick s.n, 1918, Brasil, RS, Pareci
(SP33928)
Dados não
publicados
P. eludens
Identificação
confirmada
A.A.R. de Meijer SJf-3912, 6/I/2001,
Brasil, PR, São José dos Pinhais, Roça
Velha (MBM371817)
Meijer (2006)
como P. aff.
neophlebophorus
P. eludens
Identificação
confirmada
A.A.R. de Meijer CUa-346, 28/I/1980, Meijer (2006)
como P. aff.
Reserva Natural Cambuí (MBM325664) neophlebophorus
P. exiguus
Provável P.
jamaicensis
J. Rick s.n., 1936, Brasil, RS,
Marcellino Ramos (PACA14518)
Rick (1961)
P. fallax
Não localizado
R. Singer B 110, 9/XI/1951, Brasil, RS,
Est. São Salvador (LIL)
Singer (1954) as P.
umbrinoalbidus,
Singer (1959)
P. cf. fastigiatus
(Seq. ITS disponível
para depósito)
N.K. Ishikawa & R. Vargas-Isla NKI12, Dados não
25/IV/2011, Brasil, AM, Manaus,
publicados
INPA, Campus III (INPA239970)
P. cf. fernandezianus
(Seq. ITS: JQ065028)
13/IX/2003, Brasil, RS, Passo Fundo,
Jaboticabal (RSPF330 as P. beniensis)
Dados não
publicados
P. fibulatus
Identificação
confirmada
A.A.R. de Meijer CUa-284,
14/XII/1979, Brasil, PR, São José dos
Pinhais, Reserva Natural Cambuí
(MBM324261)
Meijer (2006)
P. fibulatus
Identificação
confirmada
A.A.R. de Meijer CUa-284c,
(MBM371794)
Meijer (2006)
P. fibulatus
Identificação
confirmada
Mogi Guaçu [K(M)41525 =SP214311]
Pegler (1997)
P. fibulatus
Não localizado (ident.
confirmada duplicata)
Mogi Guaçu [SP214311 =K(M)41525]
Pegler (1997)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 244
Referências
bibliográficas
P. fibulatus
Não localizado
A.A.R. de Meijer CUa-284b, Brasil, PR, Meijer (2006)
Curitiba, São José dos Pinhais, Reserva
Natural Cambuí (MBM)
P. fluminensis
Identificação
confirmada
R. Singer B 432, 31/IX/1952, Brasil, RJ, Singer (1956) as P.
variipes, Singer
Angra dos Reis (F - holótipo)
(1959)
P. fluminensis
Identificação
confirmada
M. Capelari & R. Maziero MC212,
18/XII/1984, Brasil, SP, Cananeia, Ilha
do Cardoso, próximo Morro Três
Irmãos (SP193817)
Menolli et al.
(2010)
P. fluminensis
Identificação
confirmada
M. Capelari MC164, 23/X/1984, Brasil,
SP, Cananeia, Ilha do Cardoso
(SP194009)
Menolli et al.
(2010)
P. fluminensis
Insuficiente para
estudo
SP, Cananeia, Parque Estadual Ilha do
Cardoso, entre o Sítio dos Andradas e
Rio Santa Cruz (SP193819)
Dados não
publicados
P. cf. fuliginosus
(Seq. ITS disponível
para depósito)
04/IV/2012, Brasil, SP, São Paulo, Pq.
Est. das Fontes do Ipiranga (SP445827)
Dados não
publicados
P. fuscidulus (nom. in
herb.)
Insuficiente para
estudo
J. Rick s.n., 22/VII/1944, Brasil, RS,
São Leopoldo (PACA20926)
Dados não
publicados
P. fusconigricans
Identificação
confirmada
A.A.R. de Meijer CUa-434, 18/IV/1980, Dados não
publicados
Reserva Natural Cambuí (MBM325751)
P. glaucotinctus
Identificação
HM562147)
L.A.S. Ramos LASR64, 27/X/2009,
b)
P. glaucotinctus
Identificação
HM562157)
M. Capelari & U.C. Peixoto MC4412,
02/XII/2008, Brasil, SP, São Paulo,
(SP394384)
b)
P. glaucotinctus
Identificação
confirmada
F. Karstedt et al. FK1084, 19/II/2008,
(SP417360)
Dados não
publicados
P. glaucotinctus
Identificação
confirmada
A.A.R. de Meijer CUa-426, 26/I/1980, Meijer (2006)
como P. glaucus
P. glaucotinctus
Identificação
confirmada
(MBM371804)
P. glaucotinctus
Identificação
confirmada
A.A.R. de Meijer CUa-287b, 7/IV/1981, Meijer (2006)
como P. glaucus
Meijer (2006)
como P. glaucus
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 245
Referências
bibliográficas
P. glaucotinctus
Identificação
confirmada
A.A.R. de Meijer CUb-2085,
31/XII/1991, Brasil, PR, Curitiba,
Parque Birigui (MBM371795)
Dados não
publicados
P. glaucotinctus
Identificação
confirmada
L.K. Okino & A.M. Gugliotta 3630,
31/10/1990, Brasil, SP, Santo André,
Reserva Biológica de Paranapiacaba,
(SP233830 como P. atriavellaneus)
Dados não
publicados
P. glaucotinctus
Identificação
confirmada
V.G. Cortez 001/03 16/II/2003, Brasil,
RS, Santa Maria, Boca do Monte Passo
do Tigre (SMDB9710)
Wartchow et al.
(2006) como P.
nigrolineatus
P. glaucus
Não localizado
A.A.R. de Meijer CUa-287, Brasil, PR,
São José dos Pinhais, Reserva Natural
Cambuí (MBM)
Meijer (2006)
P. glaucus
Não localizado
Curitiba, São José dos Pinhais, Reserva
Natural Cambuí (MBM)
P. glaucus
Não localizado
A.A.R. de Meijer SJa, 9/II/1992, Brasil,
PR, São José dos Pinhais, Rua Harry
Feeken (MBM)
P. globiger
Identificação
JQ065030)
V.G. Cortez 050/04, 30/IX/2004, Brasil, Wartchow et al.
RS, Porto Alegre, Campus do
(2006)
Vale/UFRGS (ICN139025)
P. globiger
Provável Amanita
I.C. Oliveira & M.F.M Guimarães 298,
25/V/1990, Brasil, PB, João Pessoa,
Campus I da UFPB (JPB17511)
Dados não
publicados
P. globiger
Provável Amanita
I.C. Oliveira 257, 08/V/1990, Brasil,
(JPB17515)
Dados não
publicados
P. globiger
Provável Amanita
I.C. Oliveira & M.F.M Guimarães 273,
01/VIII/1990, Brasil, PB, João Pessoa,
Campus I da UFPB (JPB17537)
Dados não
publicados
P. granulatus
Provável P.
glaucotinctus
Leopoldo (PACA14523)
Rick (1961)
P. harrisii
Identificação
confirmada
A.A.R. de Meijer MAg-3717, 13/I/2000, Meijer (2010)
Brasil, PR, Morretes, Parque Marumbi, como P. cf.
Caminho do Itupava (MBM371798)
puttemansii
P. harrisii
Identificação
FJ816666, Seq. LSU:
FJ816652)
(SP393709)
Menolli et al.
(2010)
P. harrisii
Identificação
disponível)
J.J.S. Oliveira & A.V. Costa JJSO267,
06/XI/2010, Brasil, SP, Santo André,
Reserva Biológica de Paranapiacaba,
Trilha 4 (SP445849)
Dados não
publicados
Stijve & de Meijer
(1993) como
Pluteus sp. (aff.
glaucus), Meijer
(2006)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 246
Referências
bibliográficas
P. harrisii
Insuficiente para
estudo
L.H. Rosa & R.O. Morais PERD 090,
16/II/2000, Brasil, MG, Marliéria,
(SP307731)
Rosa & Capelari
(2009)
P. harrisii
Identificação
confirmada
16/II/2000, Brasil, MG, Marliéria,
(SP307732)
Rosa & Capelari
(2009)
P. harrisii
Identificação
confirmada
M. Capelari & V.L.R. Bononi MC10,
25/IV/1986, Brasil, SP, São Miguel
Arcanjo, Parque Estadual Carlos
Botelho (SP194165)
Pegler (1997),
Menolli et al.
(2010)
P. harrisii
Identificação
confirmada
M. Capelari et al. MC3282, 11/IV/1990, Menolli et al.
Brasil, SP, Brasil, Santo André, Reserva (2010)
Biológica de Paranapiacaba (SP233625)
P. harrisii
Identificação
FJ816654, Seq. LSU:
FJ816644)
Menolli Jr. et al. NMJ122, 22/III/2007, Menolli et al.
Brasil, SP, Santo André, Reserva
(2010)
P. harrisii
Identificação
HM562164 as P.
puttemansii)
(SP393698 – holótipo de P.
puttemansii)
P. harrisii
Identificação
FJ816658)
Menolli Jr. et al. NMJ132, 24/V/2007,
Menolli et al.
(2010) como
Biológica de Paranapiacaba (SP393693) Pluteus sp.
P. haywardii
Insuficiente para
estudo
07/XII/1999, Brasil, MG, Marliéria,
(SP307733)
Rosa & Capelari
(2009)
P. haywardii
Insuficiente para
estudo
M. Capelari & V.L.R. Bononi MC18,
25/IV/1986, Brasil, SP, São Miguel
Arcanjo, Parque Estadual Carlos
Botelho (SP194164)
Pegler (1997)
P. cf. haywardii
Não localizado
A.A.R. de Meijer CUa-1420, Brasil, PR, Meijer (2006)
Cambuí (MBM)
P. hispidulus
Provável P. yungensis
Salvador (PACA14528)
Rick (1961)
P. hispidulus
Rick (1961)
P. hispidulus
Entolomataceae
J. Rick s.n., 1932, Brasil, RS
(BPI839783)
Dados não
publicados
P. hispidulus
(FH00301668)
Dados não
publicados
Menolli &
Capelari (2010) as
P. puttemansii,
b)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 247
Referências
bibliográficas
P. hispidulussimilis sp.
nov.
Identificação
disponível para
depósito)
Brasil, RN, Baía Formosa, Reserva
publicados
Particular do Patrimônio Natural Mata
Estrela (SP417459 – holótipo)
P. homolae
Identificação
confirmada
A.A.R. de Meijer JSb-3259, 06/II/1996,
Brasil, PR, Jundiaí do Sul, Fazenda Pau
d'Alho (MBM371826)
P. homolae
Identificação
confirmada
publicados
P. hylaeicola
Identificação
confirmada
10/VIII/2003, Brasil, PR, Antonina,
(MBM371802)
Meijer (2006)
como P.
eucryphiae
P. hylaeicola
Não localizado
R. Singer B 4201, 05/VI/1966, Brasil,
Pará, Belém (BAFC - holótipo)
Singer (1989)
P. hylaeicola
Não localizado
R. Singer B 11273 (B12373 em Singer
Singer (1989)
1989), 12/VII/1978, Brasil, AM, Estrada
Manaus-Itacoatiara, 30 Km N of
Manaus, Embrapa (INPA82460 parátipo)
P. iguazuensis
Identificação
disponível para
depósito)
N.K. Ishikawa & R. Vargas-Isla NKI10, Dados não
25/IV/2011, Brasil, AM, Manaus,
publicados
INPA, Campus III – V-8 (INPA239968)
P. iguazuensis
Indisponível para
estudo
Brasil, RS, Frederico Westphalen
(URM80716)
Drechsler-Santos
et al. (2007)
P. jamaicensis
Insuficiente para
estudo
Jaru, margem direita do Rio Ji-Paraná
(SP212088)
Dados não
publicados
P. karstedtianus sp. nov. Identificação
disponível para
depósito)
Ponte Velha (SP445828 – holótipo)
Dados não
publicados
P. karstedtianus sp. nov. Identificação
disponível para
depósito)
Dados não
27/IV/2006, Brasil, SP, São Paulo,
publicados
Parque Estadual da Cantareira, Trilha da
Cachoeira (SP417456)
P. laetifrons var.
laetifrons
Insuficiente para
estudo
P. laetifrons var. floridae Não localizado
Dados não
publicados
Velha, propriedade de Pichorim
(MBM371796)
Meijer (2006)
A.A.R. de Meijer MAh-1670, Brasil,
PR, Piraquara, Mananciais da Serra
(MBM)
Meijer (2006)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 248
Referências
bibliográficas
P. leptonia
Entoloma s.l
Rick s.n., 1936, Brasil, RS, São
Leopoldo (PACA14525 -holótipo de P.
leptonia)
Rick (1961)
P. longistriatus
Identificação
confirmada
A.A. Carvalho Jr. 20/02, 08/XI/2002,
Brasil, RJ, Rio de Janeiro, Jardim
Botânico (RB419297)
Dados não
publicados
P. longistriatus
Identificação
confirmada na
duplicata
São Paulo, Parque Estadual das Fontes
como P. pluvialis
do Ipiranga [SP214538 = K(M)41609
como P. pluvialis]
P. longistriatus
Identificação
confirmada
como P. pluvialis
do Ipiranga [K(M)41609 = SP214538
como P. pluvialis]
P. longistriatus
Identificação
confirmada
como P. pluvialis
do Ipiranga [SP214475 = K(M)41610
como P. pluvialis]
P. longistriatus
Identificação
confirmada
F. Karstedt et al. FK679, 20/VI/2006,
das Fontes do Ipiranga, Trilha da
Nascente (SP416768)
Dados não
publicados
P. longistriatus
Identificação
confirmada
Menolli &
Capelari. (2010)
P. longistriatus
Identificação
HM562158)
Menolli &
Capelari (2010),
b)
P. longistriatus
Identificação
confirmada
Iguaçu, Trilha do Hidrômetro
(SP416766)
P. longistriatus
Provável P.
longistriatus
(FH00301670)
Dados não
publicados
P. longistriatus
Identificação
confirmada
L.A.S. Ramos & F. Karstedt LASR25,
31/VIII/2008, Brasil, SP, São Paulo,
(SP393721)
Menolli &
Capelari (2010)
P. longistriatus
Identificação
HM562149)
Menolli &
Capelari (2010),
b)
P. longistriatus
Identificação
HM562172)
25/VII/2006, Brasil, SP, São Paulo,
b)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 249
Referências
bibliográficas
P. longistriatus
Identificação
confirmada
M. Capelari et al. MC4437,
10/XII/2008, Brasil, SP, Palestina,
Fazenda Boa Vista (SP416767)
Dados não
publicados
P. longistriatus
Identificação
confirmada
M. Capelari & J.J.S. Oliveira MC4676,
(SP417464)
Dados não
publicados
P. leoninus
Provável P. conizatus
(FH00301669)
Dados não
publicados
P. maculosipes
Identificação
confirmada
Iguaçu, Trilha do Poço Preto
(SP417463)
P. microsporus
Não localizado
26/IX/1992, Brasil, SP, Águas da Prata
(SP250534)
Dados não
publicados
P. minutus
Insuficiente para
estudo
Singer B 11119, 6/VI/1978 (7/VI/1973
em Singer 1989), Brasil, BA, Itabuna
(INPA82435)
Singer (1989)
P. nanus
Provável P. sapiicola
J. Rick s.n., 29/II/1944, Brasil, RS, São
Rick (1961)
P. nanus
J. Rick s.n., 1936, Brasil, RS, Marcelino Dados não
Ramos (FH00301671)
publicados
P. necopinatus sp. nov.
Identificação
disponível para
depósito)
Ponte Velha (SP445830 – holótipo)
P. neochrysaegis sp.
nov.
Identificação
confirmada
publicados
Reserva Natural Cambuí (MBM325711
- holótipo)
P. neochrysaegis sp.
nov.
Identificação
confirmada
A.A.R. de Meijer CUa-442b,
27/V/1980, Brasil, PR, São José dos
(MBM325712)
Dados não
publicados
P. paraensis
Não localizado
R. Singer B 4238, 07/VI/1966, Brasil,
Pará, Belém (BAFC - holótipo)
Singer (1973)
P. patricius
Insuficiente para
estudo
R. Singer B 10709, 26/II/1978), Brasil,
AM, Manaus, Ponta Negra
(INPA77201)
Dados não
publicados
P. paucicystidiatus sp.
nov.
Identificação
HM562173)
Brasil, SP, Iporanga, PETAR, Núcleo
Ouro Grosso (SP394383 – holótipo)
b) como Pluteus
sp. IV
P. pellitus
Provável P. petasatus
Rick (1961)
Dados não
publicados
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 250
Referências
bibliográficas
P. pellitus
Rick (1961)
P. pellitus
(FH00301672)
Dados não
publicados
P. pellitus
Seção Hispidoderma
com pleurocistidios
pigmentados
(FH00301673)
Dados não
publicados
P. petasatus
Identificação
confirmada
R. Singer B111, Brasil, RS, São
Salvador (LIL como P. viscidulus; =
FH00301676)
Singer (1954,
1959) como P.
viscidulus
P. phlebophorus
Provável P.
tucumanus
Rick (1961)
P. phlebophorus
J. Rick s.n., 28/I/1944, Brasil, RS, São
Rick (1961)
P. cf. polycystis
Insuficiente para
estudo
(MBM324245)
Meijer (2006)
P. cf. polycystis
Insuficiente para
estudo
A.A.R. de Meijer CUb-2562, 5/III/1993, Meijer (2006)
(MBM371831)
P. cf. polycystis
Não localizado
Cambuí (MBM)
P. cf. polycystis
Não localizado
A.A.R. de Meijer CUb-2596, Brasil, PR, Meijer (2006)
Curitiba, Parque Barigui (MBM)
P. princeps (nom. in
herb.)
Provável P.
angustisporus
Meio (BPI770893)
Dados não
publicados
P. pulverulentus var.
pseudonanus
Não localizado
A.A.R. de Meijer CUa-443, Brasil, PR,
Cambuí (MBM)
Meijer (2006)
P. pulverulentus var.
pseudonanus
Não localizado
A.A.R. de Meijer SJa, 9/II/1992, Brasil,
PR, São José dos Pinhais, Rua Harry
Feeken
Meijer (2006)
P. riberaltensis var.
conquistensis
Identificação
HM562162; LSU:
FJ816648)
24/I/2008, Brasil, SP, São Paulo, Parque
Estadual das Fontes do Ipiranga
(SP393704)
Menolli &
Capelari (2010),
b)
P. rimosellus
Identificação
confirmada
A.A.R. de Meijer CUa-417, 1/IV/1980, Meijer (2006)
como P. cubensis
P. rimosellus
Identificação
confirmada
20/IV/2003, Brasil, PR, Antonina,
(MBM371803)
Meijer (2006)
como P. cubensis
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 251
Referências
bibliográficas
P. rimosellus
Identificação
confirmada
R. Singer B 433, 30/IX/1952), Brasil,
RJ, Angra dos Reis (F - holótipo de P.
subfibrillosus)
Singer (1956,
1959) como P.
subfibrillosus
P. rimosoaffinis
Identificação
HM562145)
A.M. Gugliotta et al. PEFI07/2009,
19/VIII/2009, Brasil, SP, São Paulo,
(SP394379)
b) como Pluteus
sp. III
P. rimosoaffinis
Identificação
confirmada
Mogi-Guaçu [SP214395 = K(M)41611]
P. rimosoaffinis
Identificação
confirmada na
duplicata
Mogi-Guaçu [K(M)41611 = SP214395]
P. rimosoaffinis
Identificação
confirmada
Eiten & Goodland 6313, 25/VI/1965,
Brasil, SP, São Paulo, Itapecerica da
Serra (SP106867 como P.
diptychocystis)
Pegler (1997)
como P.
diptychocystis
(SP106367)
P. rimosoaffinis
Identificação
disponível para
depósito)
M. Capelari & J.J.S. Oliveira MC4607,
Dados não
publicados
P. rimosoaffinis
Identificação
confirmada
23/X/2003, Brasil, RS, Passo Fundo,
Jaboticabal (RSPF327 como P. fallax)
Dados não
publicados
P. rimosoaffinis
Identificação
confirmada
R. Singer B11111, 6/VI/1978, Brasil,
BA, Itabuna (INPA82432)
Dados não
publicados
Não localizado
A.A.R. de Meijer CUb-2552b, Brasil,
PR, Curitiba, Parque Barigui
Meijer (2006)
como P. cf.
rimosoaffinis
Não localizado
A.A.R. de Meijer MAg, 11/IV1992,
Meijer (2006)
Brasil, PR, Morretes, Rio Nhundiaquara
(MBM)
P. riograndensis
Identificação
confirmada
A.A.R. de Meijer CUa-208, 3/II/1979,
Meijer (2006)
como P. cf.
Reserva Natural Cambuí (MBM371811) fluminensis
P. riograndensis
Identificação
confirmada
20/II/1979, Brasil, PR, São José dos
(MBM324153)
P. riograndensis
Identificação
confirmada
A.A.R. de Meijer CUa-208c, 21/I/1980, Meijer (2006)
como P. cf.
Reserva Natural Cambuí (MBM324154) fluminensis
P. riograndensis
Identificação
confirmada
A.A.R. de Meijer CUa-405, 22/III/1980, Dados não
publicados
Meijer (2006)
como P. cf.
fluminensis
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 252
Referências
bibliográficas
P. riograndensis
Identificação
confirmada
A.A.R. de Meijer CUi-2196,
18/III/1992, Brasil, PR, Curitiba,
Bosque João Paulo II (MBM371823)
Meijer (2006)
como P. cf.
rimosoaffinis
P. riograndensis
Identificação
confirmada
A.A.R. de Meijer GUf-4480,
07/IX/2009, Brasil, PR, Guaraqueçaba,
Tagaçaba, Reserva Natural Serra Itaqui
(MBM371800)
Dados não
publicados
P. riograndensis
Insuficiente para
estudo
D.L. Bordignon & J.C. Budke,
02/XI/2002, Brasil, RS, Santa Maria,
Morro do Elefante (SMDB9711 como
P. beniensis)
Wartchow et al.
(2006) como P.
beniensis
P. riograndensis
Identificação
confirmada
D.N. Pegler 3978, 3/II/1987, Brasil, SP,
Cananeia, Ilha do Cardoso [K(M)41608
=SP214350]
Pegler (1997)
como P.
fluminensis
P. riograndensis
Não localizado.
Identificação
confirmada na
duplicata
D.N. Pegler 3978, 3/II/1987, Brasil, SP,
Cananeia, Ilha do Cardoso [SP214350
=K(M)41608]
Pegler (1997)
como P.
fluminensis
P. riograndensis
Identificação
FJ816664, Seq. LSU:
FJ816650)
Menolli et al.
24/I/2008, Brasil, SP, São Paulo, Parque (2010) como P.
Estadual da Cantareira, Núcleo
fluminensis
P. riograndensis
Identificação
FJ816662)
engordador (SP393705)
Menolli et al.
(2010) como P.
fuligineovenosus
P. riograndensis
Identificação
disponível)
26/I/2011, Brasil, SC, Parque Nacional
Serra do Itajaí, Parque das Nascentes,
Trilha da 3ª Vargem (SP416738)
Dados não
publicados
P. riograndensis
Identificação
disponível)
J.J.S. Oliveira & M. Capelari JJSO09,
14/I/2010, Brasil, SP, Santo André,
(SP417454)
Dados não
publicados
P. riograndensis
Identificação
disponível)
J.J.S. Oliveira JJSO58, 16/III/2010,
Dados não
publicados
P. riograndensis
Identificação
JQ065029)
Jaboticabal (RSPF299 como P.
beniensis)
Dados não
publicados
P. riograndensis
Identificação
FJ816655)
(SP393710)
Menolli et al.
(2010) como P.
fluminensis
P. riograndensis
Identificação
FJ816657)
da Cantareira (SP393706)
Menolli et al.
(2010) como P.
jamaicensis
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 253
Referências
bibliográficas
P. riograndensis
Indisponível para
estudo
R. Singer B 31, 31/X/1951, Brasil, RS,
São Leopoldo (LIL-holótipo)
Singer (1954,
1956, 1959)
P. salicinus
Não localizado
MAc-5.xii.1992 (MBM)
Stijve & de Meijer
(1993) como P.
glaucus, Meijer
(2006)
P. sapiicola
Identificação
confirmada
Amazonas COa-3929, 15/I/2001, Brasil,
PR, Colombo, Embrapa Florestas
(MBM371806)
Meijer (2006)
como P.
pulverulentus var.
pseudonanus
P. sapiicola
Identificação
confirmada
A.A.R. de Meijer CUb-2552a,
5/III/1993, Brasil, PR, Curitiba, Parque
Barigui (MBM371822 como P.
pulverulentus var. pseudonanus)
Meijer (2006)
como P. cf.
rimosoaffinis
P. sapiicola
Identificação
HM562146)
b) como Pluteus
sp. II
P. sapiicola
Identificação
disponível)
J.J.S. Oliveira et al JJSO303,
Dados não
08/XII/2010, Brasil, SP, Santo André,
publicados
Res. Biol. de Paranapiacaba (SP416736)
P. sapiicola
Identificação
HM562148)
21/X/2009, Brasil, SP, Santo André,
(SP394382)
b) como Pluteus
sp. II
P. sapiicola
Identificação
confirmada
Parque Estadual da Cantareira (SP)
Dados não
publicados
P. sapiicola
Identificação
confirmada
M. Capelari ? - PEFI n°052/99,
Dados não
20/I/1999, Brasil, SP, São Paulo, Parque publicados
Estadual das Fontes do Ipiranga (SP)
P. scruposus
Não localizado
R. Pilger s.n., 27/II/1899, Brasil, Mato
Grosso, Cuiabá (B - holótipo)
Hennings (1900)
P. straminellus
Nomen dubium
Rick (1961)
P. striatocystis
Identificação
confirmada
A.A.R. de Meijer FE-3300, 24/II/1996,
Brasil, PR, Fênix, Parque Estadual de
Vila Rica do Espírito Santo
(MBM371821)
Meijer (2006)
como P. cubensis
P. subfibrillosus
Não localizado
Taimbesinho (LIL)
Singer (1954 como
P. rimosellus,
1956, 1959)
P. sublaevigatus
Ident. incerta,
complexo P.
aureovenatus (ITS:
FJ816667, LSU:
FJ816653)
(SP393694)
Menolli et al.
(2010)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 254
Referências
bibliográficas
P. sublaevigatus
Identificação
confirmada
A.A.R. de Meijer MAe-2611,
15/IV/1993, Brasil, PR, Marumbi
Parque (MBM371799 como P.
chrysophlebius)
Dados não
publicados
P. sublaevigatus
Identificação
confirmada
Nova Ponte, Fazenda Caxuanã, Reserva
Borda do Rio Claro (SP416737)
Dados não
publicados
P. sublaevigatus
Identificação
confirmada
F. Karstedt & J.J.S. Oliveira FK1904,
01/III/2011, Brasil, SP, São Paulo,
(SP416743)
Dados não
publicados
P. thomsonii
Identificação
confirmada
(MBM324270)
Meijer (2006)
como P. aff.
thomsonii
P. thomsonii
Indisponível para
estudo
F. Wartchow 15/X/2001, Brasil, RS,
Santa Maria, Morro do Elefante
(SMDB9194)
Wartchow et al.
(2004)
P. umbrinoalbidus
Identificação
confirmada
Menolli Jr. et al. NMJ116, 20/III/2007,
da Cantareira (SP393712)
Menolli &
Capelari (2010)
P. umbrinoalbidus
Insuficiente para
estudo
07/XII/1999, Brasil, MG, Marliéria,
(SP307734)
Rosa & Capelari
(2009)
P. umbrinoalbidus
Indisponível para
estudo (Identificação
confirmada nos
isótipos)
São Salvador (LIL- holótipo)
Singer (1954,
1956, 1959)
P. umbrinoalbidus
Identificação
confirmada
Singer B 124, 9/XI/1951, Brasil, RS,
São Salvador (MICH-isótipo =
FH00301674- isótipo = LIL-holótipo)
Singer (1954,
1956, 1959)
P. umbrinoalbidus
Indisponível para
estudo
Singer B 124, 9/XI/1951, Brasil, RS,
São Salvador (FH00301674-isótipo =
MICH-isótipo = LIL-holótipo)
Singer (1954,
1956, 1959)
P. umbrinoalbidus
Não preservado
np (MBM)
Meijer (2006)
P. umbrosus
Provável P.
angustiporus
Meio County (BPI770905)
Dados não
publicados
P. varzeicola
Identificação
confirmada
R. Singer, Araújo, Walker & Adis;
Singer B 12397, 9/III/1983), Brasil,
AM, Iranduba, Ilha Marchantaria (F holótipo)
Singer (1989)
P. varzeicola
Insuficiente para
estudo (identificação
confirmada na
duplicata)
R. Singer, Araújo, Walker & Adis;
Singer B 12397, 9/III/1983), Brasil,
AM, Iranduba, Ilha Marchantaria
(INPA108366 - isótipo = F)
Singer (1989)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 255
Referências
bibliográficas
P. varzeicola
Insuficiente para
estudo
R. Singer B 12182, 23/V/1980), Brasil,
AM, Paraná do Janauacá near teh Lago
Castanho (F)
Singer (1989)
P. velatus
Nomen dubium
Leopoldo (PACA14527 - holótipo)
Singer (1959)
como P. fibulatus,
Rick (1961)
P. velutinus
Identificação
para depósito)
PA, Belém, Ilha do Combu (SP417670) publicados
P. wehlianus
Provável Bolbitiaceae
or Strophariaceae
Rick s.n., 1934, Brasil, RS, São
Leopoldo (PACA14529 como P.
wehlianus)
P. xylophilus
Identificação
confirmada
A.A.R. de Meijer CUa-160, 3/X/1979,
Dados não
publicados
P. xylophilus
Identificação
confirmada
16/XI/1979, Brasil, PR, São José dos
(MBM324088)
Dados não
publicados
P. xylophilus
Identificação
confirmada
20/IV/2003, Brasil, PR, Antonina,
(MBM341341)
Meijer (2006,
2009)
P. xylophilus
Identificação
confirmada
A.A.R. de Meijer SJf-4023, 26/II/2001,
Velha (MBM341338)
Meijer (2006,
2009)
P. xylophilus
Identificação
confirmada
A.A.R. de Meijer SJf-4086, 28/V/2001,
Velha (MBM371816)
Meijer (2006,
2009)
P. xylophilus
Identificação
confirmada
A.I. Milanez s.n., 15/IV/1972, Brasil,
SP, Piracicaba (SP112156)
Pegler (1997)
P. xylophilus
Identificação
confirmada
Menolli et al.
(2010)
P. xylophilus
Identificação
confirmada
G. Robledo et al. s.n., 09/XI/2011,
Brasil, SP, Mogi Guaçu, Reserva
Biológica e Estação Experimental de
Mogi Guaçu, Gleba A (SP417741)
Dados não
publicados
P. xylophilus
Identificação
confirmada
G. Guzmán 22986, 10/XI/1982, Brasil,
SP, São Paulo, Parque Estadual das
Fontes do Ipiranga (SP177702)
Grandi et al.
(1984), Menolli et
al. (2010)
P. xylophilus
Identificação
confirmada
cervinus)
Rick (1961) como
P. cervinus
Rick (1961) como
P. cervinus
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 256
Referências
bibliográficas
P. xylophilus
Identificação
confirmada
cervinus)
Rick (1961) como
P. cervinus
P. xylophilus
Não Localizado
D.N. Pegler 4010, 05/II/1987, Brasil,
SP, Cananeia, Ilha do Cardoso
[SP214323 = K(M)41523]
Pegler (1997)
P. xylophilus
Identificação
confirmada
D.N. Pegler 3712, 16/I/1987, Brasil, SP, Pegler (1997),
Menolli et al.
do Ipiranga [SP214539 = K(M)41522]
(2010)
P. xylophilus
Identificação
confirmada na
duplicata
do Ipiranga [K(M)41522 = SP214539]
P. xylophilus
Identificação
confirmada
L.C. Abreu 283, 05/VIII/1975, Brasil,
SP, São Paulo, Parque Estadual das
Fontes do Ipiranga (SP128034)
Grandi et al.
(1984) e Pegler
(1997) como
Volvariella bakeri,
Menolli &
Capelari (2008)
como Pluteus sp.
P. xylophilus
Identificação
confirmada
Menolli et al.
(2010)
P. xylophilus
Identificação
disponível)
M. Capelari & P.O. Ventura MC4655,
Dados não
05/I/2012, Brasil, SP, São Paulo, Parque publicados
Estadual da Cantareira, Núcleo
P. xylophilus
Identificação
HM562163)
Menolli Jr. et al. NMJ138, 30/VI/2007,
Menolli &
Capelari (2010),
b)
P. xylophilus
Identificação
confirmada
Menolli Jr. & F. Karstedt NMJ150,
(SP393702)
Menolli et al.
(2010)
P. xylophilus
Identificação
FJ816659, Seq. LSU:
FJ816645)
Menolli Jr. et al. NMJ143, 30/VI/2007,
(SP393701)
Menolli et al.
(2010)
P. xylophilus
Identificação
confirmada
M.S. Rother & B.M.A. Severo
Jaboticabal (RSPF294)
Dados não
publicados
P. xylophilus
Identificação
confirmada
leg. Steffen s.n., det. J. Rick, 1934,
Brasil, RS, São Leopoldo (PACA14526
como P. cervinus)
Singer (1959)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 257
Referências
bibliográficas
P. cf. xylophilus
Insuficiente para
estudo
Sobestiansky 11/12, 2001, Brasil, Rio
Grande do Sul, Nova Petrópolis,
29º22'S, 51º08’W (M M191 41
Sobestiansky
(2005)
Pluteus sp.
Insuficiente para
estudo
Dados não
Amazonas COa-3628, 25/V/1999,
publicados
(MBM371824)
Pluteus sp.
Insuficiente para
estudo
Velha (MBM371815)
Meijer (2006)
como P. cubensis
Pluteus sp.
Insuficiente para
estudo
22/II/1993, Brasil, PR, São José dos
(MBM)
Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer CUh-2479,
23/II/1993, Brasil, PR, Curitiba,
Uberaba (MBM371829)
Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
publicados
Pluteus sp.
Insuficiente para
estudo
17/V/1980, Brasil, PR, São José dos
(MBM325745)
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer CUb-2549, 5/III/1993, Dados não
publicados
(MBM371832)
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer JSb-3258, 06/II/1996,
Brasil, PR, Jundiaí do Sul, Fazenda Pau
d'Alho (MBM371808)
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer CUa-332, 20/I/1980, Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer CUa-341, 25/I/1980, Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer JSa-3253, 04/II/1996,
Brasil, PR, Jundiaí do Sul, Fazenda
Monte Verde (MBM371830)
Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer JSa-3252, 04/II/1996,
Brasil, PR, Jundiaí do Sul, Fazenda
Monte Verde (MBM371807)
Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
A.A.R. de Meijer ALAa-3210,
19/I/1995, Brasil, PR, Altonia, Estrada
do Amendoin, 2 Km ao norte do Jardim
Paredão (MBM371828)
Dados não
publicados
Dados não
publicados
Dados não
publicados
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 258
Referências
bibliográficas
Pluteus sp.
Insuficiente para
estudo
Dados não
Amazonas COa-3930, 15/I/2001, Brasil, publicados
PR, Colombo, Embrapa Florestas
(MBM371797)
Pluteus sp.
Insuficiente para
identificação (Seção
Celulloderma)
A. Batista s.n., 26/II/1980, Brasil, RS,
São Francisco de Paulo, Fl. Nac. do
IBDF (ICN6833)
Pereira (1984)
Pluteus sp.
Insuficiente para
identificação
A. Batista s.n., 07/V/1980, Brasil, RS,
São Francisco de Paulo, Fl. Nac. do
IBDF (ICN6689)
Dados não
publicados
Pluteus sp.
Insuficiente para
estudo
Bionda & Raber, 19/I/1996, Brasil, RS,
Passo Fundo (RSPF124)
Dados não
publicados
Pluteus sp.
Complexo P.
aureovenatus (Seq.
ITS: HM562160)
Brasil, SP, São Paulo, Iporanga,
PETAR, Núcleo Ouro Grosso
(SP394388)
b)
Pluteus sp.
Complexo P.
aureovenatus (Seq.
ITS para depósito)
F. Karstedt FK1602, Brasil, SC, Gaspar, Dados não
Reserva Particular do Patrimônio
publicados
Natural Figueira Branca (SP416739)
Pluteus sp.
Insuficiente para
identificação
(complexo
aureovenatus)
Dados não
publicados
Parque Estadual da Cantareira, Trilha da
Cachoeira (SP416741)
Pluteus sp.
Insuficiente para
identificação
(complexo
aureovenatus)
da Cantareira, Trilha da Cachoeira
(SP416742)
Dados não
publicados
Pluteus sp.
Insuficiente para
Pluteus)
F. Karstedt FK941, 07/XI/2007, Brasil,
SP, Santo André, Reserva Biológica de
Paranapiacaba
(SP394385)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
F. Karstedt FK2014, 24/III/2011, Brasil, Dados não
BA, Camacan, RPPN Serra Bonita
publicados
(SP417461)
Pluteus sp.
Insuficiente para
identificação
Ponte Velha (SP417460)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
F. Karstedt et al. FK916, 30/V/2007,
Dados não
publicados
Pluteus sp.
Insuficiente para
Pluteus)
F. Wartchow 13/03, 15/V/2003, Brasil,
RS, Santa Maria, Arroio do Só, Água
Boa (SMDB9610)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
F. Wartchow 12, 15/X/2001, Brasil, RS, Dados não
Santa Maria, Camobi, Morro do
publicados
Elefante (SMDB9195)
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 259
Referências
bibliográficas
Pluteus sp.
Insuficiente para
estudo
G. Guzmán & V.L.R. Bononi s.n.,
Dados não
23/XII/1983, Brasil, AM, SE de Manaus publicados
(SP193567)
Pluteus sp.
Insuficiente para
Pluteus)
J. Putzke s.n., 12/VI/1992, Brasil, RS,
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
J. Putzke s.n., 13/VI/1992, Brasil, RS,
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
(SP307735)
Rosa & Capelari
(2009)
Pluteus sp.
Insuficiente para
identificação
(SP307736)
Rosa & Capelari
(2009)
Pluteus sp.
Provável
Marasmiaceae
M.A. Neves & Mueller, Pereira 290,
24/V/2008, Brasil, RN, Natal, Parque
das Dunas (JPB40696 =
HUEFS138350)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
M.A. Souza 2170, 10/III/1985, Brasil,
(JPB9272)
Dados não
publicados
Pluteus sp.
Insuficiente para
Hispidoderma)
04/XI/2009, Brasil, SP, Santo André,
(SP394377)
Dados não
publicados
Pluteus sp.
Insuficiente para
Pluteus)
21/X/2009, Brasil, SP, Santo André,
(SP394381)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
M. Capelari et al. MC1533, 14/V/1987, Dados não
Brasil, RO, Brasil, RO, Jaru, Reserva
publicados
Biológica do Jaru, margem direita do rio
Ji-Paraná, próximo à ilha 7 de setembro
(SP214566)
Pluteus sp.
Insuficiente para
identificação
M. Capelari et al. MC1467, 13/V/1987, Dados não
Brasil, RO, Brasil, RO, Jaru, Reserva
publicados
Biológica do Jaru, margem direita do rio
Ji-Paraná (SP212184)
Pluteus sp.
Insuficiente para
identificação
Jaru, margem direita do rio Ji-Paraná
(SP212053)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
M. Capelari et al. MC1271, 04/II/1987,
Brasil, SP, Cananeia, Ilha do Cardoso
(SP211778)
Dados não
publicados
Considerações
Táxons*
‡
taxonômicas/ dados
Coleção / Origem
moleculares
Apêndice C - 260
Referências
bibliográficas
Pluteus sp.
Insuficiente para
identificação
M. Capelari et al. MC1245, 03/II/1987,
Brasil, SP, Cananeia, Ilha do Cardoso
(SP211766)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
02/X/1986, Brasil, RO, Jaru, Margem
direita do Igarapé Paraíso, mata ciliar,
10°20'S - 62°10'W (SP211607)
Capelari &
Maziero (1988)
Pluteus sp.
Insuficiente para
estudo (Seq. ITS
disponível para
depósito)
Brasil, RN, Natal, Parque Estadual
publicados
Dunas de Natal (SP445823)
Pluteus sp.
Insuficiente para
identificação
M. Shinohara s.n., 27/V/1985, Brasil,
São Paulo, Vila Sonia (SP193876)
Dados não
publicados
Pluteus sp.
Insuficiente para
Pluteus)
N.K. Ishikawa et al. NKI8, 10/V/2011,
Brasil, AM, Manaus, Reserva Ducke
(INPA239966)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
P.G. Kupper s.n., 04/II/1982, Brasil,
Campos do Jordão, Parque Estadual de
Campos do Jordão (SP193698)
Dados não
publicados
Pluteus sp.
Agaricus
P.S. Silva 028/03, 20/XII/2003, Brasil,
RS, Viamão, Parque Estadual de Itapuã
(ICN139272)
Dados não
publicados
Pluteus sp.
Cortinariaceae
(ICN139275)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
(ICN139270)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
Sobestiansky 13 (MBM337561)
Dados não
publicados
Pluteus sp.
Agaricaceae –
Provável Agaricus
RTG s.n., II/1986, Brasil, RS, Viamão
(ICN56593)
Dados não
publicados
Pluteus sp.
Indisponível para
estudo
T. Kosonen, 01/I/2009, Brasil, AM,
Manaus, Universidade Federal do
Amazonas UFAM (INPA228961)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
V.G. Cortez 065/05, 20/V/2005, Brasil,
RS, Dom Pedro de Alcântara
(ICN139143)
Dados não
publicados
Pluteus sp.
Insuficiente para
identificação
V.G. Cortez 083/05, 10/IX/2005, Brasil, Dados não
RS, Viamão, Parque Estadual de Itapuã publicados
(ICN139161)
Créditos - 261
CRÉDITO DAS IMAGENS
CAPA – Arte final: Luiz Antonio Silva Ramos.
PÁG. 56
Figura 3: Pluteus albostipitatus. (a–m) Fernanda Karstedt. (n) Alfredo Justo. (o–p) Horak &
Heinemann (1978).
PÁG. 63
Figura 6: Pluteus glaucotinctus. (a–c) Marina Capelari. (d–h). Pedro Arrillaga.
PÁG. 67
Figura 8: Pluteus longistriatus. (a, b, d, e). Fernanda Karstedt. (c). Marina Capelari.
(f). Walter J. Sundberg.
PÁG. 94
Figura 7: Pluteus crassus. (a–d) Nelson Menolli Junior.
PÁG. 98
Figura 9: Pluteus fibrillosus. (a–c). Nelson Menolli Junior
PÁG. 127
Figura 23: Pluteus riograndensis. (a–c). Nelson Menolli Junior
PÁG. 155
Figura 2: (a–c). Pluteus argentinensis – Fernanda Karstedt. (d–f). P. maculosipes – Fernanda
Karstedt. (g–i). P. velutinus – Fernanda Karstedt.
PÁG. 185
Figura 2: (a, b). Pluteus aureolatus – Fernanda Karstedt. (c, f). P. brunneocrinitus – Marina
Capelari. (d, e) P. cebolinhae – Fernanda Karstedt. (g, h). P. crinitus – Fernanda Karstedt.
(i). P. diptychocystis – Iuri Goulart Baseia. (j). P. cf. fastigiatus – Noemia Kazue Ishikawa.
(k, l). P. cf. fuliginosus – Fernanda Karstedt. (m, n). P. hispidulussimilis – Iuri Goulart Baseia.
(o–p). P. karstedtianus – Fernanda Karstedt. (q–r). P. necopinatus – Fernanda Karstedt.
PÁG. 185
Figura 14. (a–e). Pluteus paucicystidiatus. (a). Dennis E. Desjardin. (b–e). Nelson Menolli
Junior. (f). P. rimosoaffinis – Jadson José Souza de Oliveira

nelson menolli junior - Atendimento a alunos de pós

Transcrição

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