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Tetrapyrgos atrocyanea, the type species of Tetrapyrgos, and Campanella buettneri, the type species of Campanella, are redescribed and epitypified based on recently collected material from Madagascar and Príncipe, respectively, supported with morphological and LSU, ITS sequences data. Line drawings, colour photographs, and comparisons with similar taxa are provided.
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Submitted 23 May 2017, Accepted 22 June 2017, Published 2 July 2017
Corresponding Author: Dennis E. Desjardin e mail 977
The type species of Tetrapyrgos and Campanella (Basidiomycota,
Agaricales) are redescribed and epitypified
Desjardin DE1*, Perry BA2, Shay JE1, Newman DS1 and Randrianjohany E3
1Department of Biology, San Francisco State University, 1600 Holloway Ave., San Francisco, CA 94132, USA
2Department of Biology, California State University East Bay, 25800 Carlos Bee Blvd., Hayward, CA 94542, USA
3Centre National de Recherches sur l'Environnement (CNRE), BP 1739, Lab. de Microbiologie de l'Environnement (LME),
Antananarivo, Madagascar
Desjardin DE, Perry BA, Shay JE, Newman DS, Randrianjohany E 2017 The type species of Tetrapyrgos and
Campanella (Basidiomycota, Agaricales) are redescribed and epitypified. Mycosphere 8(8), 977985, Doi
Tetrapyrgos atrocyanea, the type species of Tetrapyrgos, and Campanella buettneri, the type species of Campanella,
are redescribed and epitypified based on recently collected material from Madagascar and Príncipe, respectively,
supported with morphological and LSU, ITS sequences data. Line drawings, colour photographs, and comparisons with
similar taxa are provided.
Key Words agarics Campanella buettneri fungal diversity taxonomy Tetrapyrgos atrocyanea
There has been renewed interest in clarifying the generic boundaries of Tetrapyrgos E. Horak (1987) and
Campanella Hennings (1895). The morphological distinctions between these two genera and the ways they have been
treated taxonomically by various workers were succinctly documented by Honan et al. (2015). However, they noted
that distinguishing Tetrapyrgos from Campanella was problematical because of the lack of recently collected specimens
of their type species upon which comprehensive morphological and molecular redescriptions can be based. The
holotype specimen of the type species of Tetrapyrgos (Pterospora atrocyanea Métrod) was collected in 1935 in
Madagascar, and is stored in alcohol (PC), rendering it of limited molecular phylogenetic value. The holotype specimen
Mycosphere 8(8): 977985 (2017) ISSN 2077 7019
Doi 10.5943/mycosphere/8/8/1
Copyright © Guizhou Academy of Agricultural Sciences
of the type species of Campanella (C. buettneri Henn.) was collected in 1892 in Togo, is also stored in liquid (B), and
of limited value for molecular analyses. Our recent fieldwork in Madagascar and the West African islands of São Tomé
and Príncipe (approx. 700 miles from Togo) have yielded specimens that we recognize as Tetrapyrgos atrocyanea and
Campanella buettneri, respectively. The purpose of this paper is to redescribe these two important type species, based
on morphological characteristics of these recent specimens which we designate as epitype material, and to provide ITS
and nLSU sequences to aid in distinguishing the species from others, and for future research. Determining whether
Tetrapyrgos and Campanella represent a single, morphologically variable genus is beyond the scope of this
Materials and Methods
Morphological observations
Fresh material of Tetrapyrgos atrocyanea was collected in Madagascar in Febuary 2014, and fresh material of
Campanella buettneri was collected in São Tomé and Príncipe April 2006 and 2008. Macromorphological notes and
photographs were documented soon after collection, before drying the basidiomes for transport and herbarium storage.
Colour terms and notations in parentheses are those of Kornerup & Wanscher (1978). Micromorphological analyses
were performed using dried material rehydrated in 100% ethanol followed by Melzer’s reagent, or Congo Red Solution
and 3% KOH, and documented using a Nikon Optiphot-2 compound microscope fitted with a drawing tube. Because
of the unusual tetrahedral shape of basidiospores of Tetrapyrgos, their length and width were determined by measuring
the sides of a quadrilateral into which each spore would fit. Spore statistics were calculated as: xm, the arithmetic mean
of the spore length by the spore width standard deviation) from n spores measured in a single specimen; Q, the
quotient of spore length and spore width in any one basidiospore indicated as a range of variation in n spores measured;
Qm, the mean of Q-values in a single specimen; n, the number of spores measured per specimen; s, the number of
specimens studied. Specimens are deposited in the H.D. Thiers Herbarium at San Francisco State University (SFSU).
Molecular methods
Total genomic DNA was extracted from dried material using the Extract-N-Amp Plant PCR Kit (Sigma-
Aldrich, St. Louis, MO) following the manufacturer instructions. PCR protocols followed those outlined in Perry et al.
(2007). The nuclear ribosomal internal transcribed spacer region (ITS) and ribosomal large subunit (nLSU) were
amplified using primer pairs ITS1-F/ITS4 (Gardes and Bruns 1993; White et al. 1990) and LROR/LR7 (Moncalvo et.
al. 2000), respectively. Amplification products were cleaned using the Exo-SAPit kit (Affymetrix, Santa Clara, CA),
and sent to ELIM Biopharmaceuticals (Hayward, CA) for sequencing. Resulting sequencing products were edited,
assembled, aligned and compared to top BLAST hits in Geneious 9.0 (Biomatters Ltd., Auckland, New Zealand). All
sequences generated as part of this study have been deposited in GenBank (accessions MF075136 MF075139).
Tetrapyrgos atrocyanea (Métrod) E. Horak, Sydowia 39: 102. 1987 (1986). Figs. 1, 2ae
Basionym: Pterospora atrocyanea Métrod, Prod. Fl. Mycol. Madag. 3: 140. 1949.
Holotype MADAGASCAR. Antananarivo (as Tananarive), in a pasture in a government park, 16 Dec. 1934, coll. by
R Decary, No. 1642 (PC).
Epitype hic designari MADAGASCAR. Region Analamanga, Antananarivo, Parc Botanique de Zoologique, near the
garden of Crops Wild and Relatives, S18˚55.530', E47˚31.350', elevation 1270 m, 8 Feb. 2014, coll. by DS Newman
and JE Shay, JES 216 (MF075139LSU, MF075137ITS, SFSU).
MycoBank: MB 131349; Facesoffungi number: 03427
Description of epitype material:
Pileus 525 mm diam, convex to plano-convex, centrally slightly depressed in age; margin striate; surface dull,
dry, felted; disc initially brownish grey (5C13), developing dark bluish grey tones in age, margin initially white,
becoming pinkish buff to orangish white (5A3) in age, staining dark bluish grey (20E34) in splotches. Context up to
1 mm thick, pliant, tenacious, concolorous with pileus surface, staining dark bluish grey. Lamellae adnate with a
decurrent tooth to arcuate, close (1925) with 23 series of lamellulae, moderately broad (12 mm), white, often
becoming pale orange white (5A2) in age, sometimes staining bluish grey; edges pruinose. Stipe 1036 12 mm,
central, cylindrical or narrowed downward, tough, pliant, hollow, non-insititious, arising from a flattened pad of bluish
black mycelium; surface dull, dry, felted-pruinose; apex white when young, becoming orangish white (5A2) to
brownish grey (5EF3) in age; base brownish grey (5F3) to dark bluish grey (20DE34) to bluish black (20F45).
Fig. 1 Basidiomes of Tetrapyrgos atrocyanea (Epitype, JES 216). Scale bars = 10 mm
Photographs by DS Newman
Basidiospores (8) 1011.8 (6.5) 79 µm [xm = 11.0 ± 0.63 7.9 ± 0.75 µm, Q = 1.131.77, Qm = 1.41 ±
0.27, n = 20, s = 1], tetrahedral, hyaline, inamyloid, thin-walled. Basidia 3040 78.5 µm, clavate, 4-spored, hyaline.
Basidioles subclavate to subfusoid. Pleurocystidia absent. Lamellar edge sterile. Cheilocystidia abundant, 3050 4
5 µm, irregularly cylindrical, densely diverticulate, usually with a knob-like apex; apical knob 514 3.87 µm, broadly
obtuse, smooth; diverticula 1.55 13.5 µm, cylindrical to knob-like, obtuse; hyaline, inamyloid, non-gelatinous,
thin-walled. Subhymenium non-gelatinous. Pileipellis a well-developed Rameales-structure of hyphae 3.57 µm diam,
irregularly cylindrical, sometimes forked, densely diverticulate; terminal cells repent to erect, similar to the
cheilocystidia, typically with a knob-like apex; apical knob 516 4.59 µm, broadly obtuse, smooth; diverticula 1.5
5 1.53.5 µm, cylindrical to knob-like, obtuse; hyaline, inamyloid, non-gelatinous. Pileus trama of loosely interwoven
hyphae 25 µm diam, cylindrical to irregular in outline, hyaline, inamyloid, strongly gelatinous. Lamellar trama regular,
hyphae 25 µm diam, cylindrical to irregular in outline, hyaline, inamyloid, strongly gelatinous. Stipitipellis similar to
the pileipellis, a well-developed Rameales-structure of densely diverticulate hyphae with apical knobs; stipe cortical
hyphae 2.55 µm diam, parallel, cylindrical, hyaline, inamyloid, non-gelatinous, thin- to thick-walled; stipe medullary
hyphae 310 (12) µm diam, cylindrical, hyaline, inamyloid, non-gelatinous. Clamp connections common in all tissues.
Fig. 2 Micromorphological features of Tetrapyrgos atrocyanea (Epitype, JES 216). a. Basidiospores. b. Basidium
and basidioles. c. Cheilocystidia. d. Pileipellis terminal cells. e. Stipitipellis cells. Scale bar = 10 µm. Illustration by
DE Desjardin.
Habit, habitat and known distribution Gregarious, sometimes subcespitose, on woody debris amongst
Casuarina, Panicum, Galinsoga, Commelina and bamboo, in a botanical garden and zoological park. Madagascar.
Notes Tetrapyrgos atrocyanea (Bas. Pterospora atrocyanea Métrod) is the type species of the genus
Tetrapyrgos, established by Horak (1987) to replace the invalid Pterospora Métrod (1949) (non Pterospora Nuttall
1818, Pyrolaceae). The holotype specimen was collected in a government park in Antananarivo, Madagascar, and
comprehensively described in the protologue (Métrod 1949). The holotype, preserved in alcohol, was studied by Horak
(1968) who provided more details on micromorphology. As noted by Honan et al (2015), the documentation of
tetrahedral basidiospores, Rameales-type pileipellis and stipitipellis, diverticulate cheilocystidia with apical knobs,
bluish stains, and a stipe with white apex and bluish black base provided enough data to help circumscribe the genus
Tetrapyrgos. However, the poorly preserved holotype and absence of recently collected material prohibited generation
of molecular sequences to help distinguish the taxon from similar species, especially from T. longicystidiata A.H.
Honan, Desjardin & T.J. Baroni, newly described from neotropical populations, from T. subcinerea (Berk. & Broome)
E. Horak, described from Sri Lanka and widely distributed throughout south and southeast Asia, and from the genus
Campanella. During recent fieldwork in Madagascar, co-authors Shay and Newman encountered several specimens of
Tetrapyrgos, and one in particular (JES 216) was collected in a government park in Antananarivo, quite possibly the
same park from which the holotype originated. We herein accept this specimen as epitype material for T. atrocyanea.
The morphology of JES 216 matches quite nicely that described in the protologue (Métrod 1949) and
subsequent type study (Horak 1968), differing only slightly in basidiospore size. Métrod (1949) reported the
basidiospores as "78 2.53.5 µm, ovoid with two lateral wings 22.5 µm long" (collectively 78 6.58.5 µm),
while Horak (1968) noted the basidiospores as tetrahedral, 79 µm broad and long. Our material shows distinctly
"winged", tetrahedral basidiospores 811.8 79 µm, i.e., up to several microns longer than reported for the holotype
Pairwise comparisons of aligned, overlapping ITS sequences of the epitype of T. atrocyanea with the top ten
BLAST results indicate 99.9% similarity to the holotype specimen of T. longicystidiata (EF175544) from Puerto Rico,
and 99.6% similarity to a specimen of T. longicystidiata from Bolivia (EF175533). In the protologue of T. atrocyanea
(Métrod 1949; pg. 129, Fig. 87c) and Horak's type study (Horak 1968; pg. 528, Fig. e) they illustrated very long
cheilocystidia (4555 µm sensu Horak), the diagnostic feature of T. longicystidiata (Honan et al. 2015; pg. 109, Fig.
2d). These data suggest that the recently described new world taxon is a synonym of T. atrocyanea, described from
Campanella buettneri Henn., Bot. Jb. 22: 95. 1895. Figs. 3, 4ad
Holotype AFRICA, Togo, Bismarckburg, on wood, 1892, coll. by R. Büttner (B).
Epitype hic designari AFRICA, Príncipe island, Bom Bom resort area, N1˚41.559', E7˚24.171', 20 April 2008, coll.
by D.E Desjardin, DED 8276 (MF075138LSU, MF075136ITS, SFSU)
MycoBank: MB 208007; Facesoffungi number: 03428
Description of epitype material:
Pileus 512 mm diam, auriform, asymmetrically convex to campanulate in profile, hemispherical to
subchordate in face view, margin undulate, sometimes cleft; surface rugulose to pustulose, tessellate, dull, dry, white
overall, drying white to pale cream, unstaining. Context thin, rubbery-pliant, white. Lamellae adnate to point of
attachment, remote with many cross-lamellae, costate-reticulate, intervenose, relatively narrow, white, drying dark
cream, unstaining. Stipe absent; with or without a poorly developed pseudostipe extending from the margin, usually in
the cleft, or eccentrically from the pileus surface. Odor and taste not distinctive.
Basidiospores 8.610.2 6.07.2 µm [xm = 9.7 ± 0.46 6.6 ± 0.34 µm, Q = 1.41.6, Qm = 1.48 ± 0.04, n = 26, s = 1],
broadly ellipsoid, rarely approaching subglobose, lacking bulges, smooth, hyaline, inamyloid, thin-walled. Basidia 28
32 78 µm, clavate, 4-spored. Basidioles clavate to subclavate, seldom subfusoid. Cheilocystidia not differentiated.
Hymenial cystidia common on sides and edges of lamellae, 3848 912 µm, fusoid to ventricose, hyaline, thin-walled
or with wall up to 0.5 µm thick (not metuloids). Pileipellis a well-developed Rameales-structure; hyphae 35 µm diam,
cylindrical to irregular in outline, dendrohyphidia-like, densely diverticulate, hyaline, inamyloid, thin-walled, non-
gelatinous; diverticula 212 12 µm, cylindrical to narrowly conical, mostly obtuse, hyaline, thin-walled. Pileus trama
of loosely interwoven hyphae 24 (5) µm diam, cylindrical, non-diverticulate, hyaline, inamyloid, embedded in a
strongly gelatinous matrix. Lamellar trama regular, hyphae 2.55 µm diam, hyaline, inamyloid, gelatinous. Clamp
connections common in all tissues.
Fig. 3 Basidiomes of Campanella buettneri (Epitype, DED 8276). Scale bar = 10 mm
Photograph by BA Perry
Habitat and known distribution Gregarious on woody debris of undetermined dicotyledonous plants. Africa
(Cameroon, Príncipe, São Tomé, Togo), Hawaii.
Additional material examined São Tomé, Parque Nacional Obo, along trail from Bom Succeso to Lagoa
Amelia, N0˚17.317', E6˚36.746', 28 April 2006 and 2 May 2006, coll. by D.E. Desjardin, material lost in transit. United
States. Hawaiian Islands, Hawai`i, Lava Tree State Park near Leilani, N19˚28'08", W154˚54'10", 30 July 2005, coll. by
Ed Lickey, TFB 12565 (TENN-F-60782).
Notes Campanella buettneri is the type species of the genus Campanella, established by Hennings (1895) from
material collected in Togo and Cameroon. We are unaware of recently collected material from these countries upon
which the generic concept of Campanella can be refined, but we have collected material from the nearby island nation
of São Tomé and Príncipe that we recognize as representing this important species, and we herein accept it as epitype
material for C. buettneri.
Fig. 4 Micromorphological features of Campanella buettneri (Epitype, DED 8276). a. Basidiospores. b. Basidium
and basidioles. c. Hymenial cystidia. d. Pileipellis hyphae. Scale bar = 10 µm. Illustration by DE Desjardin.
The Príncipe specimen matches the protologue (Hennings 1895) and differs only slightly from an analysis of
the holotype specimen studied by Singer (1975). Campanella buettneri is characterized by small, rubbery-pliant
basidiomes with a white, ear-shaped, tessellate-verrucose pileus 512 mm diam attached to the substrate by a small,
eccentric to lateral pseudostipe, costate-reticulate, intervenose, white hymenophore, broadly ellipsoid basidiospores
with mean 9.7 6.6 µm, fusoid-ventricose hymenial cystidia, a Rameales-type pileipellis, and growth on woody debris.
The protologue reported the basidiospores as subglobose, 79 µm, while Singer (1975) reported those in the holotype
specimen as ellipsoid, 8.7 6.3 µm; neither author mentioned lateral bulges on the basidiospores. The basidiospores of
the Príncipe material are mostly broadly ellipsoid with only a few tending towards subglobose, but the range in variation
encompasses the sizes reported by Hennings (1895) and Singer (1975); likewise, they lack lateral bulges. Additional
noteworthy features of the species include the distinct fusoid-ventricose, non-refractive, thin- to firm-walled hymenial
cystidia, strongly gelatinous pileus trama, a non-gelatinous pileipellis of irregular hyphae with relatively long, narrow
diverticula, and white basidiomes that do not discolor greyish green.
Singer (1975) compared C. buettneri to C. alba (Berk. & M.A. Curtis) Singer, a species described from Cuba
and rather widespread in the neotropics, indicating that the latter differed primarily in having basidiospores with lateral
bulges. Campanella alba differs also in forming few, scattered hymenial cystidia that are narrowly cylindrical (2.77.5
µm) and subcapitate to capitate, whereas in C. buettneri the hymenial cystidia are fusoid-ventricose, 912 µm diam,
and non-capitate. Pairwise comparisons of aligned, overlapping ITS sequences of C. buettneri with the top ten BLAST
results indicate closest similarity (99.5%) to a sequence determined as C. alba (DQ449943) obtained from a specimen
collected on Hawai`i island, USA. Examination of the latter specimen (TENN-F-60782) indicated morphology nearly
identical to that of the epitype specimen designated herein (albeit with fewer hymenial cystidia), and we recognize the
Hawaiian material as representing C. buettneri, not C. alba.
São Tomé and Príncipe: We thank Dr. Robert C. Drewes (California Academy of Sciences) who continues to
initiate, coordinate and lead multiorganism biotic surveys on São Tomé and Príncipe; Eng. Arlindo de Ceita Carvalho,
Director General of the Ministry of Environment, Victor Bonfim, Salvador Sousa Pontes and Danilo Barbero for
permission to collect and export specimens for study. We are indebted to Société de Conservation et Développement
for logistics and housing support, especially the wonderful staffs of Omali Lodge and Bom Bom Island. We are grateful
for the support and cooperation of Bastien Loloumb of Zuntabawe and Faustino Oliviera, former Director of the
botanical garden at Bom Sucesso. We were assisted in the field by Jose Ramos Maria Vital Pires on Príncipe and by
Quintino Quade Cabral, Martinho Nascimiento and Jose Clara on São Tomé. For continuing support, we are most
grateful to Ned Seligman, Quintino Quade Cabral and Roberta dos Santos of STePUP. We are grateful to the College
of Science and Engineering at San Francisco State University for partial funding to support travel to São Tomé and
Príncipe, and to the G. Lindsay Field Research Fund of the California Academy of Sciences (CAS) for financially
supporting the expedition in 2006 and the Hagey Research Venture Fund (CAS) in 2008.
Madagascar: We are grateful to Rokiman Letsara of the California Academy of Science, Madagascar
Biodiversity Research Station for his invaluable efforts in planning and facilitating field work. We are thankful for the
knowledge of local mycologist Emile Randrianjohany and his colleagues at the Centre National de Recherches sur
l'Environnement (CNRE) for their valuable field assistance.
We thank the Department of Biology, San Francisco State University, Mycological Society of San Francisco, Sonoma
County Mycological Association, and the generous donors from a Kickstarter campaign who partially funded fieldwork.
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... Tetrapyrgos was proposed by Horak (1987) to replace Pterospora Métrod (1949:129) that has priority as the name of a genus of angiosperm within Ericaceae. Tetrapyrgos atrocyanea (Métrod) E. Horak (1987:102), originally described from Madagascar and the type species of the genus, was recently recollected in the type locality and the specimen was established as the epitype (Desjardin et al. 2017) from which an ITS barcode sequence was provided. Horak (1983Horak ( , 1987 recognized 15 species as belonging to the genus, including taxa formerly classified in Campanella Henn. ...
... or ITS. The phylogenetic placement of T. atrocyanea among the Tetrapyrgos sampling is displayed in trees which resulted from phylogenetic analyses shown in this paper, including the ITS sequence generated from the epitype (Desjardin et al. 2017). Tetrapyrgos atrocyanea is reported for the first time from the Brazilian Amazon. ...
... Tetrapyrgos atrocyanea is reported for the first time from the Brazilian Amazon. Also, based on the phylogenetic trees and on morphological evaluation, T. longicystidiata is herein synonymized under T. atrocyanea as suggested in Desjardin et al. (2017). Detailed descriptions of macro-and micromorphological structures are provided for the new species, along with information regarding the ecology and location of the specimens, taxonomic discussion, color photographs and line-drawings. ...
Tetrapyrgos is characterized by small basidiomata which have a mostly whitish pileus, a well-defined central or eccentric, mostly dark greyish stipe, a non-or very weakly gelatinous pileipellis and pileus tramal tissues; the pileipellis is composed of Rameales-like structures, and the basidiospores are distinctly tetrahedral. Members of this genus are commonly found on dead leaves and woody debris on the floor of tropical and subtropical forests. Recent studies suggest changes in the circumscription of Tetrapyrgos and Campanella Still poorly documented, to date, only about 18 species have been assigned to Tetrapyrgos. Through integrative taxonomy, we describe six new species from the Brazilian Amazon, based on evidence from morphological and ITS data. Tetrapyrgos longicystidiata is formally synonymized in T. atrocyanea.
... Notes:-tetrapyrgos atrocyanea is the type species of the genus tetrapyrgos Horak. It was described from Madagascar by Métrod (1949) and an epitype was recently designated by Desjardin et al. (2017), together with a detailed description and molecular sequences. tetrapyrgos atrocyanea has a wide distribution, in addition to Madagascar it is known from Benin (Piepenbring et al. 2020), Bolivia, Brazil (Komura et al. 2020), Costa Rica, and Puerto Rico (Desjardin et al. 2017;Honan et al. 2015). ...
... It was described from Madagascar by Métrod (1949) and an epitype was recently designated by Desjardin et al. (2017), together with a detailed description and molecular sequences. tetrapyrgos atrocyanea has a wide distribution, in addition to Madagascar it is known from Benin (Piepenbring et al. 2020), Bolivia, Brazil (Komura et al. 2020), Costa Rica, and Puerto Rico (Desjardin et al. 2017;Honan et al. 2015). Often, tetrapyrgos specimens in the tropics and subtropics were referred to as tetrapyrgos nigripes (Fries 1838: 383) E. Horak (Horak 1983;Piepenbring et al. 2018;Redhead 1989;Singer 1973). ...
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Mushrooms and allies belong to the Agaricomycetes lineage of Basidiomycota. A total of 260 species, belonging in 109 genera, 51 families and 13 orders have been reported from São Tomé and Príncipe between 1851 and 2020, of which 66 were described as new species. They range in body forms from agarics and boletes to polyporoid, clavarioid, coralloid, thelephoroid, stereoid, corticioid, hydnoid, cantherelloid, gasteroid, and jelly fungi. The vast majority are saprotrophs, a small number are plant pathogens, and a rare few may be ectomycorrhizal. Sixty species, 23%, can be classified putative endemics. The current state of knowledge of the Agaricomycetes from the nation is based on fewer than ten expeditions in the past 170 years and represents only a snapshot of the actual diversity that is likely present.
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São Tomé e Príncipe é arquipélago localizado no Golfo da Guiné, África Ocidental. A ilha de São Tomé é um hotspot de biodiversidade, apresentando elevadas taxas de endemismo. Estas taxas de endemismo são maioritariamente reportadas para plantas vasculares, aves, repteis, anfíbios e organismos marinhos. A diversidade fúngica da ilha de São Tomé tem vindo a ser estudada desde 1851, somando já cerca de 300 espécies de macrofungos reportados até 2019. No entanto, o conhecimento micológico é escasso e disperso, pelo que a necessidade de um estudo aprofundado é premente. O objetivo deste trabalho foi compilar todas as informações acerca da diversidade macrofúngica da ilha de São Tomé e adicionar novas informações obtidas em inventários micológicos in situ. Através de análise bibliográfica e missões de campo, foi possível construir uma lista de espécies de macrofungos e avaliar o seu potencial comestível e medicinal. Foi também possível avaliar a etnomicologia local, que se resume a quatro nomes comuns: Utu, Utu-sandjá, Cloçon-son e Ntuda renda. Tentou-se fazer a correspondência entre estes nomes comuns e os nomes científicos. O conhecimento dos cogumelos comestíveis e medicinais de São Tomé será uma ferramenta extremamente útil para o futuro das populações desta ilha, uma vez que o projeto Tesouros d’Obô pretende utilizar este conhecimento para estabelecer pequenas unidades de produção de cogumelos em comunidades-alvo. Palavras-chave: São Tomé; Fungos; Cogumelos; África; Funga; Micologia; Cogumelos comestíveis; Cogumelos medicinais; África Ocidental; Diversidade fúngica São Tomé e Príncipe is an archipelago situated on the Gulf of Guinea, West Africa. São Tomé’s island is a biodiversity hotspot, with reported high endemism rates. These endemism rates are mainly reported for vascular plants, birds, reptiles, amphibians, and marine organims. São Tomé’s island’s fungal diversity has been studied since 1851, summing up to around 300 macrofungi species reported until 2019. However, the mycological knowledge is scarce and dispersed, therefore the need for a deeper study is pressing. The objective of this work was to compile all of the information related to the macrofungal diversity of São Tomé’s Island and add new information obtained in in situ mycological inventories. Through the bibliographic analysis and the field missions, it was possible to build a species list and evaluate the edibility and medicinal potential of those species. It was also possible to evaluate the local ethnomycology, that sums up to four common names: Utu, Utu-sandjá, Cloçon-son e Ntuda renda. It was attempted to establish the link between these common names and the respective scientific names. The knowledge of São Tomé’s edible and medicinal mushrooms will be an extremely useful tool for the future of this island’s populations because the Tesouros d’Obô project is aiming to use this knowledge to establish small mushroom production units in the target communities. Keywords: São Tomé; Fungi; Mushrooms; Africa; Funga; Mycology; Edible mushrooms; Medicinal mushrooms; West Africa; Fungal diversity
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The Basidiomycota constitutes a major phylum of the kingdom Fungi and is second in species numbers to the Ascomycota. The present work provides an overview of all validly published, currently used basidiomycete genera to date in a single document. An outline of all genera of Basidiomycota is provided, which includes 1928 currently used genera names, with 1263 synonyms, which are distributed in 241 families, 68 orders, 18 classes and four subphyla. We provide brief notes for each accepted genus including information on classification, number of accepted species, type species, life mode, habitat, distribution, and sequence information. Furthermore, three phylogenetic analyses with combined LSU, SSU, 5.8s, rpb1, rpb2, and ef1 datasets for the subphyla Agaricomycotina, Pucciniomycotina and Ustilaginomycotina are conducted, respectively. Divergence time estimates are provided to the family level with 632 species from 62 orders, 168 families and 605 genera. Our study indicates that the divergence times of the subphyla in Basidiomycota are 406–430 Mya, classes are 211–383 Mya, and orders are 99–323 Mya, which are largely consistent with previous studies. In this study, all phylogenetically supported families were dated, with the families of Agaricomycotina diverging from 27–178 Mya, Pucciniomycotina from 85–222 Mya, and Ustilaginomycotina from 79–177 Mya. Divergence times as additional criterion in ranking provide additional evidence to resolve taxonomic problems in the Basidiomycota taxonomic system, and also provide a better understanding of their phylogeny and evolution.
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Tetrapyrgos (Basidiomycota, Agaricales) is recognized as a genus characterized by basidiomes with a central or eccentric, well-developed stipe with a black to bluish black stipe base arising from a basal pad or disc; hyaline, inamyloid, distinctly tetrahedral basidiospores; non-gelatinous or weakly gelatinous pileipellis and pileus tramal tissues; and apically bulbous cheilocystidia with diverticulate central axis. It is distinguished from Campanella, which has basidiomes that are sessile or with a pseudostipe that lacks black or bluish black pigmentation, and does not arise from a pad or basal disc; basidiospores ellipsoid or bulging slightly on one side; strongly gelatinous pileipellis and pileus tramal tissues; and cheilocystidia typically with central portion of axis non-diverticulate. Analyses and taxonomic status of 16 species placed in Tetrapyrgos (and their reported synonyms) are provided, based on examinations of 80 recently collected specimens and 85 exsiccata. Nine species are recognized here in Tetrapyrgos, supported by morphological and molecular data, including two new species, T. longicystidiata and T. parvispora. The remaining 7 species are recognized as belonging in Campanella or are of uncertain taxonomic placement. Comprehensive descriptions for all accepted species, type studies, and phylogenetic inferences derived from ITS analyses are provided.
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Phylogenetic relationships of mushrooms and their relatives within the order Agaricales were addressed by using nuclear large subunit ribosomal DNA sequences. Approximately 900 bases of the 5' end of the nucleus-encoded large subunit RNA gene were sequenced for 154 selected taxa representing most families within the Agaricales. Several phylogenetic methods were used, including weighted and equally weighted parsimony (MP), maximum likelihood (ML), and distance methods (NJ). The starting tree for branch swapping in the ML analyses was the tree with the highest ML score among previously produced MP and NJ trees. A high degree of consensus was observed between phylogenetic estimates obtained through MP and ML. NJ trees differed according to the distance model that was used; however, all NJ trees still supported most of the same terminal groupings as the MP and ML trees did. NJ trees were always significantly suboptimal when evaluated against the best MP and ML trees, by both parsimony and likelihood tests. Our analyses suggest that weighted MP and ML provide the best estimates of Agaricales phylogeny. Similar support was observed between bootstrapping and jackknifing methods for evaluation of tree robustness. Phylogenetic analyses revealed many groups of agaricoid fungi that are supported by moderate to high bootstrap or jackknife values or are consistent with morphology-based classification schemes. Analyses also support separate placement of the boletes and russules, which are basal to the main core group of gilled mushrooms (the Agaricineae of Singer). Examples of monophyletic groups include the families Amanitaceae, Coprinaceae (excluding Coprinus comatus and subfamily Panaeolideae), Agaricaceae (excluding the Cystodermateae), and Strophariaceae pro parte (Stropharia, Pholiota, and Hypholoma); the mycorrhizal species of Tricholoma (including Leucopaxillus, also mycorrhizal); Mycena and Resinomycena; Termitomyces, Podabrella, and Lyophyllum; and Pleurotus with Hohenbuehelia. Several groups revealed by these data to be nonmonophyletic include the families Tricholomataceae, Cortinariaceae, and Hygrophoraceae and the genera Clitocybe, Omphalina, and Marasmius. This study provides a framework for future systematics studies in the Agaricales and suggestions for analyzing large molecular data sets.
We have designed two taxon-selective primers for the internal transcribed spacer (ITS) region in the nuclear ribosomal repeat unit. These primers, ITS1-F and ITS4-B, were intended to be specific to fungi and basidiomycetes, respectively. We have tested the specificity of these primers against 13 species of ascomycetes, 14 of basidiomycetes, and 15 of plants. Our results showed that ITS4-B, when paired with either a ‘universal’ primer ITS1 or the fungal-specific primer ITS1-F, efficiently amplified DNA from all basidiomycetes and discriminated against ascomycete DNAs. The results with plants were not as clearcut. The ITS1-F/ITS4-B primer pair produced a small amount of PCR product for certain plant species, but the quantity was in most cases less than that produced by the ‘universal’ ITS primers. However, under conditions where both plant and fungal DNAs were present, the fungal DNA was amplified to the apparent exclusion of plant DNA. ITS1-F/ITS4-B preferential amplification was shown to be particularly useful for detection and analysis of the basidiomycete component in ectomycorrhizae and in rust-infected tissues. These primers can be used to study the structure of ectomycorrhizal communities or the distribution of rusts on alternate hosts.
Partial sequences of nuLSU rDNA were obtained to investigate the phylogenetic relationships of Pyronemataceae, the largest and least studied family of Pezizales. The dataset includes sequences for 162 species from 51 genera of Pyronemataceae, and 39 species from an additional 13 families of Pezizales. Parsimony, ML, and Bayesian analyses suggest that Pyronemataceae is not monophyletic as it is currently circumscribed. Ascodesmidaceae is nested within Pyronemataceae, and several pyronemataceous taxa are resolved outside the family. Glaziellaceae forms the sister group to Pyronemataceae in ML analyses, but this relationship, as well as those of Pyronemataceae to the other members of the lineage, are not resolved with support. Fourteen clades of pyronemataceous taxa are well supported and/or present in all recovered trees. Several pyronemataceous genera are suggested to be non-monophyletic, including Anthracobia, Cheilymenia, Geopyxis, Humaria, Lasiobolidium, Neottiella, Octospora, Pulvinula, Stephensia, Tricharina, and Trichophaea. Cleistothecial and truffle or truffle-like ascomata forms appear to have evolved independently multiple times within Pyronemataceae. Results of these analyses do not support previous classifications of Pyronemataceae, and suggest that morphological characters traditionally used to segregate the family into subfamilial groups are not phylogenetically informative above the genus level.