Phytotaxa 183 (4): 239253

www.mapress.com/phytotaxa/
Copyright 2014 Magnolia Press

Article

ISSN 1179-3155 (print edition)

PHYTOTAXA

ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.183.4.3

A new species and new records of gasteroid fungi (Basidiomycota) from Central
Amazonia, Brazil
TIARA S. CABRAL1, BIANCA D. B. DA SILVA2, NOEMIA K. ISHIKAWA3, DONIS S. ALFREDO4, RICARDO
BRAGA-NETO5, CHARLES R. CLEMENT6 & IURI G. BASEIA7
Programa de Ps-graduao em Gentica, Conservao e Biologia Evolutiva; Instituto Nacional de Pesquisas da AmazniaINPA; Av.
Andr Arajo, 2936Petrpolis; Manaus, Amazonas, 69067-375 Brazil. Email: ttiara@gmail.com
2
Programa de Ps-graduao em Sistemtica e Evoluo; Universidade Federal do Rio Grande do Norte; Natal, Rio Grande do Norte,
59072-970 Brazil. Email: biancadeni@yahoo.com.br
3
Coordenao de Biodiversidade; INPA; Manaus, Amazonas, 69067-375 Brazil. Email: noemia.kazue@gmail.com
4
Programa de Ps-graduao em Sistemtica e Evoluo; Universidade Federal do Rio Grande do Norte; Natal, Rio Grande do Norte,
59072-970 Brazil. Email: donis.alfredo@yahoo.com.br
5
Centro de Referncia em Informao Ambiental (CRIA); Av. Romeu Trtima, 388; Campinas, So Paulo 13084-791, Brazil. Email:
saci@cria.org.br
6
Coordenao de Tecnologia e Inovao; INPA; Manaus, Amazonas, 69067-375 Brazil. Email: cclement@inpa.gov.br
7
Departamento de Botnica e Zoologia; Universidade Federal do Rio Grande do Norte; Natal, Rio Grande do Norte 59072-970, Brazil.
Email: iuribaseia@gmail.com
1

Abstract
A new species, Geastrum inpaense, is described morphologically and molecularly. Geastrum lloydianum, G. schweinitzii,
Phallus merulinus and Staheliomyces cinctus are reported here as new records for Central Amazonia. In addition, Mutinus
fleischeri is reported as a new record for the Americas and Phallus atrovolvatus, as a new record for Brazil.

Introduction
In the past few years, studies involving gasteroid fungi have intensified in Brazil, resulting in new species, new
occurrence data of known species, and first records of known species for several biomes (Hennings 1904, Capelari
& Maziero 1988, Baseia & Milanez 2002, Baseia et al. 2003, Baseia & Calonge 2005, 2006, Leite et al. 2007, Silva
et al. 2007, Cortez et al. 2008, Fazolino et al. 2008, Gurgel et al. 2008, Trierveiler-Pereira et al. 2009, Fazolino et
al. 2010, Ottoni et al. 2010, Leite et al. 2011, Trierveiler-Pereira et al. 2011, Cruz et al. 2012, Alves & Cortez 2013,
Alfredo et al. 2012a, 2012b, 2014, Silva et al. 2013, Alfredo & Baseia 2014, Cabral et al. 2014). The northern region
of Brazil harbors most of the megadiverse Amazon rainforest (MMA 2002), the richest assemblage of plant species
and the largest pool of tropical carbon on Earth (ter Steege et al. 2013). The Amazon River basin drains approximately
6.9 million km2, about 40% of South America, and about two thirds of it are in Brazil, with the rest in Bolivia, Peru,
Ecuador, Colombia, Venezuela, Guyana, Suriname and French Guiana. Approximately 80% of the basin is forested,
with both open and dense evergreen (ombrophilous) forests on the uplands (terra firme) and in the major floodplains
(vrzea). The rest is a mosaic of savannas (cerrado), white-sand scrub (campinas), open swamps (pntanos) and forested
stream swamps (igap), and other less common ecosystems (Veloso et al. 1991). Although numerous species of fungi
have been found in Amazonia, studies involving gasteroid mycobiota are still emerging, with few species recorded for
this region to date.

This study contributes to the expansion of knowledge about gasteroid fungi in Central Amazonia, and is based on
both molecular and morphological data. We describe a new species, Geastrum inpaense sp. nov., and register Mutinus
fleischeri Penzig (1899:137) as a first record for the American continent, Phallus atrovolvatus Kreisel & Calonge
(2005:6) as a new record for Brazil and Geastrum lloydianum Rick (1906:27), G. schweinitzii (Berk. & M.A. Curtis)
Zeller (1948:649), Phallus merulinus (Berk.) Cooke (1882:57) and Staheliomyces cinctus E. Fischer (1921:142) as
new records for Central Amazonia.
Accepted by Genevieve Gates: 10 Sep. 2014; published: 24 Oct. 2014

239

Materials and Methods

MorphologyThe specimens were collected in the state of Amazonas in the municipalities of Codajs and Manaus
(Campus I and III, Tropical Forestry Experiment Station, Adolpho Ducke Forest Reserve and Adolpho Ducke Botanical
Garden of the National Institute for Amazonian ResearchINPA). The new species was described morphologically from
fresh material according to Calonge et al. (2004) and the new records identified from Surez & Wright (1996), Kreisel
(1996) and Calonge et al. (2005). Microscopically, we measured 20 of each of the necessary taxonomic structures for
species identification using a Leica EZ4 stereomicroscope and Olympus BX42 optical microscope (OM). In addition,
scanning electron microscopy (SEM) Philips XL30 was used, following the usual methodology in this type of study
(Cortez et al. 2008, Silva et al. 2011). For the color analyses we followed Kppers (1979). Following Braga-Neto et
al. (2013) we used decimal degrees as a standard form of geographic coordinates. The specimens were deposited in the
INPA herbarium and duplicates in the UFRN herbarium. Primary data about specimens analyzed are openly and freely
available through the Brazilian Virtual Herbarium of Plants and Fungi (http://inct.splink.org.br/index).

Molecular methodsWe conducted a phylogenetic analysis to compare the morphologically close species
Geastrum inpaense sp. nov. and Geastrum albonigrum Calonge & M. Mata (2004:332). DNA extraction was performed
according to Cabral et al. (2012). The nuclear ribosomal ITS and nuc-LSU regions, and mitochondrial atp6 region
were amplified with previously described primers and protocols (Vigalys & Hester 1990, Kretzer & Bruns 1999,
Hosaka 2009). After purification of PCR fragments with ExoSAP-IT (Affymetrix Inc.), sequencing was performed
with BigDye Terminator Cycle Sequencing Ready Reaction Kit version 3.1 with the same primers.

Sequences of Geastrum and Myriostoma specimens from Kasuya et al. (2012), Silva et al. (2013), and those
generated in this study (Table 01), were aligned with Clustal X 2.1 (Larkin et al. 2007) and then manually edited with
BioEdit (Hall 1999); ambiguously aligned regions were excluded. A distance matrix was calculated only with the ITS
region using Kimura-2-parameters substitution model with MEGA (Tamura et al. 2007) to evaluate the divergence
between species. We conducted Maximum Parsimony (MP) and Bayesian phylogenetic analyses with concatenated
ITS, nuc-LSU and atp6 sequences. For Maximum Parsimony we used PAUP* (Swofford 1998), in which trees were
calculated using a heuristic search with the TBR algorithm for branch-swapping; the initial tree was obtained by
stepwise addition of random additional sequences repeated 100 times, and bootstrap of 1000 replicates. The Bayesian
Analysis was conducted with MrBayes v.3.1.2 (Huelsenbeck & Ronquist 2001), with substitution models chosen by
MrModeltest (Nylander 2004) for each partition. This analysis consisted of two different runs with four incrementally
heated simultaneous MCMC simulations over 2 million generations. Trees were sampled every 1000 generations and
parts of the trees were discarded as a burn-in stage to estimate posterior probabilities, observing the average standard
deviation of split frequency values. Trees that resulted from both analyses were edited with FigTree (http://tree.bio.
ed.ac.uk/software/figtree/).

Results
Morphological and phylogenetic analysesA total of 12 species was identified morphologically, of which one,
Geastrum inpaense sp. nov., is new to science. Mutinus fleischeri is a new record for the Americas, and Phallus
atrovolvatus is a new record for Brazil. New records for Central Amazonia are Geastrum lloydianum, G. schweinitzii,
Phallus merulinus and Staheliomyces cinctus.

The phylogenetic analysis included 28 specimens and 1899 characters, of which 587 from the atp6 region, 523
from the ITS region, and 789 from the nuc-LSU region. For phylogenetic analyses, 446 of the 1899 characters were
informative for Maximum Parsimony (MP) analysis, which resulted in a most parsimonious tree with 1725 steps (CI
= 0.523, RI = 0.564, RC = 0.295). In the Bayesian analysis, the first 4900 trees were discarded in a burn-in period;
the consensus tree was calculated with the remaining 15100. The trees generated by MP and Bayesian analysis have
slightly different topologies in the position of the clusters, but both show Geastrum inpaense sp. nov. as a sister group
to G. albonigrum with high support values (Figure 01). These data are also available in TreeBASE under ID 15201.
The distance matrix calculated with ITS shows null divergence values among the three Geastrum inpaense sp. nov.
specimens, and 0.0534 between these and G. albonigrum. In addition, null divergence values were also found between G.
pectinatum Persoon (1801: 132) (JN845116) and G. triplex Junghuhn (1840: 287) (JN845168), and 0.0034 divergence
between G. fimbriatum Fries (1829: 16) (JN845094) and G. sessile (Sowerby) Pouzar (1971: 95) (JN845123). The
low levels of divergence between these species may result from errors of morphological identifications and/or
240 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.

manipulation of sequences, and do not invalidate the divergence values found between Geastrum inpaense sp. nov. and
G. albonigrum. Thus, the divergence values based on the ITS region between these species suggest two molecularly
distinct evolutionary units.

FIGURE 1. Phylogenetic tree of Geastrum used to position G. inpaense within the genus constructed with Bayesian analysis, with
posterior probabilities values on nodes. Herbarium vouchers follow the taxa name.

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 241

Discussion
Taxonomy
Geastraceae Corda, Icones fungorum hucusque cognitorum 5: 25 (1842)
Geastrum inpaense T.S. Cabral, B.D.B. Silva & I.G. Baseia, sp. nov. (Fig. 2, 3 and 4)
MB 807545.
Diagnosis:Unexpanded basidiomata depressed-globose. Expanded basidiomata with a hairy mycelial layer.
Endoperidial sac subglobose, sessile, apophysis absent, peristome fibrillose to sulcate, delimited and lighter than
endoperidium. Basidiospores globose, verrucose, with flattened end columnar ornamentation.

Holotype:BRAZIL. Amazonas: Manaus, Instituto Nacional de Pesquisas da Amaznia-Campus I (Latitude:
3.094986, Longitude:059.986811, Datum: WGS84), growing on decomposed wood, 23 April 2011, leg. TS Cabral
(INPA 239990 holotype); Amazonas: Manaus, Jardim Botnico Adolpho Ducke, on wood, 22 January 2013, leg. TS
Cabral (INPA 255834 paratype). Genbank accession number for ITS sequence: KJ127023.
Etymology:Inpaense, refers to the place where the holotype was found.

Unexpanded basidiomata depressed-globose, 1014 mm diam. Expanded basidiomata 3240 mm wide (including
rays) 1014 mm tall, rhizomorphs present. Exoperidium saccate splitting into 5 to 9 rays, not hygroscopic. Mycelial
layer brownish (A50M20C30), with an external hirsute layer of aggregate hairs up to 0.5 mm long, and an internal cottony
layer, incrusted with debris and detaching irregularly from the external layer. Fibrous layer pale yellow (N00Y20M00),
coriaceous, with longitudinal fissures joining at the center of a convex base. Fleshy layer brown (Y90M80C70), up to 1
mm thick. Endoperidial sac 1516 mm diam., 1214 mm tall including peristome, subglobose, dark brown (Y90M80C80),
sessile. Apophysis absent. Peristome fibrillose to irregularly sulcate, mammiform, distinctly delimited and darker than
the endoperidium. Mature gleba black (N99C00A00). No subiculum noted.

Basidiospores globose 2.63.8 m (including ornamentation), brown in 5% KOH, ornamented with long, sparse
and flattened end columns measuring 0.20.3 m. Basidia clavate to irregularly clavate, with three to four sterigmata.
Capillitial hyphae 2.56.4 m diam., sinuous, rugose, walls up to 1 m thick, brownish in 5% KOH. Mycelial layer
made up of sinuous thin-walled hyphae 25 m diam., with lumen, yellow to hyaline in 5% KOH. Fibrous layer
composed of thick-walled sinuous to straight hyphae with short ramifications and clamp connections, narrow lumen,
hyaline in 5% KOH. Fleshy layer made up of subglobose hyphae 2270 m diam., yellowish to brown in 5% KOH.

Additional specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus I, 12 April 2011, Oliveira, GL
& Ishikawa, NK (INPA 240011); Amazonas: Manaus, INPA Campus I, 23 April 2011, Cabral TS 14 (INPA 239994);
Cabral TS 13 (INPA 239993); Amazonas: Manaus, INPA Campus I, 2 May 2011, Cabral TS 15 (INPA 239995);
Amazonas: Manaus, INPA Campus I, 9 May 2011, Cabral TS 17 (INPA 239997).

Habitat:Found on rotten wood. Specimens were collected in fragments of secondary upland forests, a
successional type of lowland ombrophilous dense forest.

Comments:Geastrum inpaense can be distinguished from G. albonigrum by the length of the hairs of the
hirsute layer (up to 0.6 mm long), delimited peristome and smaller basidiospores (up to 3.8 m diam.). Phylogenetic
analysis indicates that the specimens are a distinct species. Geastrum inpaense resembles G. hirsutum Baseia &
Calonge (2006: 302) in the lignicolous habitat, hirsute mycelial layer and delimited peristome, but in G. hirsutum the
rhizomorphs are absent, the spores have verrucose ornamentation and it grows on a subiculum. Geastrum javanicum
Lveill (1846:161) and G. schweinitzii are also lignicolous, however, they do not have a hirsute mycelial layer. In
G. javanicum the mycelial layer peels off and G. schweinitzii has smaller basidiomata. Molecular and morphological
analyses support the establishment of G. inpaense as a new species.

New Records
Geastrum lageniforme Vittad. [as Geaster], Monograph Lyc.: 16 (1842).
Description:Trierveiler-Pereira et al. (2011: 580)

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forests, a
successional type of lowland ombrophilous dense forest. It is known from Europe (Dissing & Lange 1962, Pegler et
al. 1995), Africa, North America, Asia (Dissing & Lange 1962).
242 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.

FIGURE2.Geastrum inpaensemacrostructures. Expanded basidiomata (a, b, c, d), scale bars correspond to 20 mm; peristome (e, f),
scale bars correspond to 5 mm; hairs of mycelial layer indicated by arrows (g, h), scale bars correspond to 10 mm and 1 mm. Photographs
by T.S. Cabral.

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 243

FIGURE3.Geastrum inpaensemicrostructures. Endoperidium surface (a), spore ornamentation (b), capillitial hyphae with (c) and
without (d) amorphous substance.

Specimens examined:BRAZIL. Amazonas: Manaus, INPA, Campus III (Latitude:3.092820, Longitude:
59.994046, Datum: WGS84), 18 January 2011, Cotegano, C.A.A. et al. 1 (INPA 240008).
Geastrum lloydianum Rick [as Geaster], Brotria, sr. bot. 5: 27 (1906)
Description:Trierveiler-Pereira et al. (2011: 581)

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forest, a
successional type of lowland ombrophilous dense forest. It is known from Tropical America (Ponce de Len 1968).

This is a first record for the state of Amazonas, Brazil.

Specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus I (Latitude:3.094986, Longitude:
59.986811, Datum: WGS84), 19 April 2011, Cabral TS 9 (INPA 239989), 19 April 2011, Cabral TS 1 (INPA239981);
23 April 2011, Cabral TS 12 (INPA 239992); 15 May 2011, Cabral TS 23 (INPA 240004); 18 May 2010, Ishikawa NK
16 (INPA240017).

Geastrum saccatum Fr., Syst. mycol. (Lundae) 3(1): 16 (1829) (Fig. 5b)
Description:Leite et al. (2011:390).

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forest, a
successional type of lowland ombrophilous dense forest. It is known from Asia (Bottomley 1948, Cunningham 1944,
Liu 1984); Africa, North America, South America, Europe, Oceania (Bottomley 1948, Cunningham 1944, Dissing &
Lange 1962, Liu 1984). This is a first record for the state of Amazonas, Brazil.

Specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus I (Latitude:3.094986, Longitude:
59.986811, Datum: WGS84), 10 May 2011, Cabral TS 16 (INPA239996).

244 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.

FIGURE 4.Geastrum inpaenseschematic drawing. Mycelial layer hyphae (a), bar = 10 m; fibrous layerhyphae (b), bar = 10 m; fleshy
layer hyphae (c), bar = 20 m; spores (d), bar = 10 m; basidia (e), bar = 10 m. Illustration by D.S. Alfredo.

Geastrum schweinitzii (Berk. & M.A. Curtis) Zeller, Mycologia 40(6): 649 (1948) (Fig. 5c).
Description:Trierveiler-Pereira et al. (2011: 582)

Habitat and distribution:on rotten wood. Specimens were collected in fragments of old-growth and secondary
upland forests, a type of lowland ombrophilous dense forest. It is known from Africa, North America, South America,
A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 245

Asia, Oceania (Ponce de Len 1968). Although it is a common species in the tropics, this is the first record for
Amazonia.

Specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus I (Latitude:3.094986, Longitude:
59.986811, Datum: WGS84), 10 May 2011, Cabral TS 6 (INPA239986), 19 April 2011, Cabral TS 8 (INPA239988);
Cabral TS 7 (INPA 239987); 11 April 2011, Ishikawa NK 11 (INPA240006); INPA Campus III (Latitude:3.092820,
Longitude:59.994046, Datum: WGS84), 11 April 2011, Vargas-Isla R 12 (INPA 240007); Estao Experimental de
Silvicultura Tropical, 10 May 2011, Cabral TS 19 (INPA239999).

FIGURE
5.Geastraceae
species.Geastrum
triplex(d).Photographs by T.S. Cabral.

lloydianum(a),Geastrum

saccatum(b),Geastrum

schweinitzii(c),Geastrum

Geastrum triplex Jungh. [as Geaster], Tijdschr. Nat. Gesch. Physiol. 7: 287 (1840) (Fig. 5d).
Description:Trierveiler-Pereira et al. (2011: 583)

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forests, a
successional type of lowland ombrophilous dense forest. It is known from Africa, North America, South America,
Oceania (Bottomley 1948, Cunningham 1944, Dissing & Lange 1962, Liu 1984); Asia (Cunningham 1944, Dissing &
Lange 1962, Liu 1984); Europe (Bottomley 1948, Cunningham 1944, Liu 1984, Dissing & Lange 1962, Moyersoen &
Demoulin 1996, Pegler et al. 1995). This is the first record for the state of Amazonas, Brazil.
Specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus I (Latitude:3.094986, Longitude:59.986811,
Datum: WGS84), 20 November 2012, Cabral TS 33 (INPA 255832); Universidade Federal do AmazonasMini Campus
(Latitude:3.101082, Longitude:59.974573, Datum: WGS84), 22 December 2012, Cabral TS 34 (INPA 255833).
Phallaceae Corda, Icones fungorum hucusque cognitorum 5: 29 (1842)
Mutinus fleischeri Penz., Ann. Jard. Bot. Buitenzorg, suppl. 16: 137 (1899) (Fig. 6d)
Description:Li et al. (2002: 126)

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forest, a
successional type of lowland ombrophilous dense forests. It is known from Asia (Li et al. 2002). Although it is unlikely
246 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.

to find a species with such a disjunct distribution, based on the morphological data we prefer to maintain it as Mutinus
fleischeri until further molecular analyses are carried out to compare Amazonian Mutinus species. This is the first
record for the American continent.

Specimens examined:BRAZIL. Amazonas: Manaus, INPA Campus III (Latitude:3.092820, Longitude:
59.994046, Datum: WGS84), 03 December 2010, Karstedt, F. et al. (INPA 240012).

FIGURE 6.Phallaceae species.Phallus merulinus(a),Phallus atrovolvatus(b),Phallus indusiatus(c),Mutinus fleischeri(d),Staheliomyces

cinctus(e). Scale bars correspond to 20 mm. Photographs (a), (b) and (d) by N.K.Ishikawa, photographs (c) and (e) by T.S. Cabral.

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 247

Phallus atrovolvatus Kreisel & Calonge, Bull. Soc. Micol. Madrid 29: 58 (2005) (Fig. 6b)
Description:Cheype (2010: 56)

Habitat and distribution:On sandy soil with rotten bamboo leaves. Specimens were collected in fragments of
secondary upland forests, a successional type of lowland ombrophilous dense forests. It is known from Central and
South America (Calonge et al. 2005, Cheype 2010). This is the first record for Brazil.

Specimens examined:BRAZIL. Amazonas: Manaus, Tarum, 15 November 2010, Ishikawa, NK 15 (INPA
240016).
Phallus indusiatus Vent.: Pers., Syn. Meth. Fung. 244 (1801) (Fig. 6c)
Synonyms: Dictyophora phalloidea Desv., J. Bot. (Paris), 2, p. 92 (1809).
Dictyophora tahitensis (Schltdl.) E. Fich., Jahrb. Knigl. Bot. Gart. Berlin, 4, p. 37 (1886).
Phallus callichrous (Mller) Lloyd, Mycol. Writ., 2, p. 6 (1907).

Description:Baseia et al. (2006: 89).

Habitat and distribution:On soil, next to roots of aa (Euterpe precatoria Mart., Palmae). Specimens were
collected in fragments of secondary floodplain forest, a successional type of alluvial ombrophilous dense forest. It
is known from Asia, Africa, North America, South America and Oceania (Bottomley 1948, Cunningham 1944, Liu
1984). This is the first record for the state of Amazonas, Brazil.

Specimens examined:BRAZIL. Amazonas: Codajs (Latitude:3.78291, Longitude:62.02186, Datum:
WGS84), 12 December 2012, Cabral TS 29 (INPA 255830).
Phallus merulinus (Berk.) Cooke, Grevillea 11(no. 58): 57 (1882) (Fig. 6a)
Synonyms: Dictyophora irpicina Pat., Bull. Soc. mycol. Fr., 14, p. 190 (1898).
Phallus irpicinus (Pat.) Lloyd, Mycol. Writ., 2 (26), p. 331 (1907).

Description:Cheype et al. (2010: 54)

Habitat and distribution:Specimens were collected on sandy soil in fragments of secondary upland forests, a
successional type of lowland ombrophilous dense forest. It is known from South America (Cheype 2010) and Asia
(Liu 1984). This is the first record for Central Amazonia and the second record for Brazil; the first record was from the
southern state of Paran as Phallus cf. merulinus (Meijer 2006).

Specimens examined:BRAZIL. Amazonas: Manaus, ASSINPA (Latitude:3.092724, Longitude:59.994434,
Datum: WGS84), 12 November 2010, Ishikawa NK (INPA240010); INPA Campus III (Latitude:3.092820, Longitude:
59.994046, Datum: WGS84), 10 November 2010, Ishikawa NK 12 (INPA240009); Puraquequara, 30 November 2010,
Vargas-Isla R. 13 (INPA 240014); Puraquequara, 29 November 2010, Vargas-Isla R. 15 (INPA240015).
Staheliomyces cinctus E. Fisch., Mitt. naturf. Ges. Bern: 142 (1921) [1920] (Fig. 6e)
Description:Leite et al. (2007: 123)

Habitat and distribution:On rotten wood (Pouteria spp., Sapotaceae). Specimens were collected in fragments
of secondary floodplain forests, a successional type of alluvial ombrophilous dense forests. It is known from Central
and South America (Calonge et al. 2005, Cheype 2010). This is the first record for Central Amazonia.

Specimens examined:BRAZIL. Amazonas: Codajs (Latitude:3.78580, Longitude:62.01911, Datum:
WGS84), 12 December 2012, Cabral TS 30 (INPA 255831).
Agaricaceae Chevall., Flore Gnrale des Environs de Paris 1: 121 (1826)
Morganella fuliginea (Berk. & M.A. Curtis) Kreisel & Dring, Feddes Repert. 74: 113 (1967). (Fig. 7)
Description:Surez and Wright (1996: 657)

Habitat and distribution:Growing gregariously and cespitose on dead wood. Specimens were collected in
lowland ombrophilous dense forests, both in old-growth and secondary upland forests. It is known from Africa (Dring
1964) and South America (Surez and Wright 1996). This is the first record for the state of Amazonas, Brazil.
248 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.


Additional specimens examined:BRAZIL. Manaus: Reserva Florestal Adolpho Ducke (Latitude:02.92803,
Longitude:059.96860, Datum: WGS84), 28 January 2008, R. Braga-Neto, GF 38 (INPA 239561); 30 October 2012,
T.S. Cabral TSC 26 (INPA 255838); 27 (INPA 255839).

FIGURE 7.Morganella fuliginea. Expanded basidiomata (a); exoperidium sphaerocyst chains (b); spores in 5% KOH (c) and under SEM
(d). Photograph by R.B. Neto.

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 249

TABLE 1. Genbank accession numbers for the Geastrum and Myriostoma species used in this phylogenetic analysis to
position G. inpaense within the genus. Sequences generated in this study are in bold.
Species

Herbarium Voucher

ITS

LSU

atp6

Geastrum albonigrum

UFRN-Fungos 1989

KJ127026

KJ127019

KJ127015

Geastrum coronatum

S:F-34813

JN845092

JN845210

JN845335

Geastrum entomophilum

UFRN-Fungos 1233

KJ127032

KJ127022

Geastrum fimbriatum

L:837179

JN845093

JN845211

JN845336

Geastrum fimbriatum

TENN:61511

JN845094

JN845212

JN845337

Geastrum hirsutum

UFRN-Fungos 1214

KJ127029

JQ683662

JQ683670

Geastrum hungaricum

TNS:TKG-GE-90502

JN845096

JN845214

JN845339

Geastrum inpaense

INPA239990

KJ127023

KJ127017

KJ127013

Geastrum inpaense

INPA255834

KJ127024

KJ127018

KJ127014

Geastrum inpaense

INPA240011

KJ127025

Geastrum javanicum

UFRN-Fungos 1215

KJ127031

JQ683663

KJ127016

Geastrum javanicum

TNS:TKG-GE-90902

JN845100

JN845218

JN845342

Geastrum minimum

K:154623

JN845105

JN845223

JN845347

Geastrum mirabile

TNS:KH-JPN10-711

JN845108

JN845226

JN845350

Geastrum mirabile

TNS:KH-JPN10-675

JN845106

JN845224

JN845348

Geastrum morganii

UFRN-Fungos 1794

KJ127028

KJ127020

Geastrum parvistriatum

JCZ 285

JN943161

JN939571

Geastrum pectinatum

S:F-46074

JN845116

JN845234

JN845358

Geastrum quadrifidum

TNS:TKG-GE-91002

JN845118

JN845236

JN845360

Geastrum saccatum

TENN:61141

JN845120

JN845238

JN845362

Geastrum schweinitzii

UFRN-Fungos 1741

KJ127030

JQ683664

JQ683671

Geastrum sessile

TENN:39858

JN845123

JN845241

JN845365

Geastrum setiferum

UFRN-Fungos 803

KJ127027

KJ127021

Geastrum striatum

S:F-46048

JN845124

JN845242

JN845366

Geastrum triplex

TENN:61723

JN845168

JN845292

JN845399

Geastrum velutinum

PDD:REB2886

JN845173

JN845297

JN845404

Myriostoma coliforme

TNS:TKG-GE-50801

JN845203

JN845328

Myriostoma coliforme

QCNE:M3353

JN845327

JN845434

Acknowledgements
The authors thank CAPES (PNADB program), CNPq (473422/2012-3) and FAPEAM (3137/2012) for financial
support, CAPES and CNPq for scholarships to authors 1, 2 and 4, the PPBio and INCT-CENBAM programs, and
Victor Py-Daniel and Ruby Vargas-Isla for collecting support. The authors also thank Doriane Picano Rodrigues for
coordination of the Applied Evolution Laboratory at the Federal University of Amazonas, where molecular analyses
were carried out.

References
Alfredo, D.S., Leite, A.G., Braga-Neto, R. & Baseia, I.G. (2012a) Two new Morganella species from the Brazilian Amazon rainforest.
Mycosphere 3(1): 6671.

http://dx.doi.org/10.5943/mycosphere/3/1/8
Alfredo, D.S., Leite, A.G., Braga-Neto, R., Cortez, V.G. & Baseia, I.G. (2012b) Scleroderma minutisporum, a new earthball from the
Amazon rainforest. Mycosphere 3(3): 294299.

250 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.


http://dx.doi.org/10.5943/mycosphere/3/3/4
Alfredo, D.S., Accioly & T., Baseia, I.G. (2014) Morganella arenicola, a new species record from North and Northeast Brasil. Turkish
Journal of Botany 38: 595599.

http://dx.doi.org/10.3906/bot-1307-68
Alfredo, D.S. & Baseia, I.G. (2014) Morganella nuda, a new puffball (Agaricaceae, Basidiomycota) in the upland forests of the Brazilian
semi-arid region. Nova Hedwigia 98: 459466.

http://dx.doi.org/10.1127/0029-5035/2014/0164
Alves, C.R. & Cortez, V.G. (2013) Morganella sulcatostoma sp. nov. (Agaricales, Basidiomycota) from Paran State, Brazil. Nova
Hedwigia 96: 409417.

http://dx.doi.org/10.1127/0029-5035/2013/0078
Baseia, I.G. & Calonge, F.D. (2005) Asero floriformis, a new phalloid with a sunflower shaped receptacle. Mycotaxon 92: 169172.
Baseia, I.G. & Calonge, F.D. (2006) Geastrum hirsutum: a new earthstar fungus with a hairy exoperidium. Mycotaxon 95: 301304.
Baseia, I.G. & Milanez, A.I. (2002) Geastrum setiferum (Gasteromycetes): A new species with a setose endoperidium. Mycotaxon 84:
135139.
Baseia, I.G., Gilbertoni, T.B. & Maia, L.C. (2003) Phallus pygmaeus, a new minute species from a Brazilian tropical rain forest. Mycotaxon
85: 7779.
Bottomley A.M. (1948) Gasteromycetes of South Africa. Bothalia 4: 473810.
Braga-Neto, R., Giovanni, R., Pezzini, F.F., Canhos, D.A.L., Marino, A., Souza, S. & Maia, L.C. (2013) Spatial data for fungal specimens:
retrospective georeferencing and practical recommendations for mycologists. Mycotaxon 125: 289301.

http://dx.doi.org/10.5248/125.289
Cabral, T.S., Marinho, P., Goto, B.T. & Baseia, I.G. (2012) Abrachium, a new genus in the Clathraceae, and Itajahya reassessed. Mycotaxon
119: 419429.

http://dx.doi.org/10.5248/119.419
Cabral, T.S., Silva, B.D.B., Marinho, P. & Baseia, I.G. (2014) Geastrum rusticum (Geastraceae, Basidiomycota), a new earthstar fungus
in the Brazilian Atlantic rainforesta molecular analysis. Nova Hedwigia 98(12): 265272.

http://dx.doi.org/10.1127/0029-5035/2013/0158
Calonge, F.D. & Mata, M. (2004) A new species of Geastrum from Costa Rica and Mxico. Boletn de la Sociedad Micolgica de Madrid
28: 331335.
Calonge, F.D., Kreisel, H. & Mata, M. (2005) Phallus atrovolvatus sp. nov. from Costa Rica. Boletn de la Sociedad Micolgica de Madrid
29: 58.
Capelari, M. & Maziero, R. (1988) Fungos macroscpicos do Estado de Rondnia Regio dos Rios Jaru e Ji-Paran. Hoehnea 15: 28
36.
Chevallier, F.F. (1826) Flore Gnrale des Environs de Paris. Chez Ferra Jeune, Paris, 648 pp.
Cheype, J.L. (2010) Phallaceae et Clathrus rcolets en Guyane franaise. Bulletin Mycologique et Botanique Dauphin-Savoie 197:
5166.
Cooke, M.C. (1882) Australian fungi. Grevillea 11(58): 5765.
Corda, A.C.J. (1842) Icones fungorum hucusque cognitorum [...] Tomus V. Apud J. G. Calve, Prague, 92 pp.
Cortez, V.G., Baseia, I.G., Guerrero, R.T. & Silveira, R.M.B. (2008) Two sequestrate cortinarioid fungi from Rio Grande do Sul, Brasil.
Hoehnea 35: 511516.
Cruz, R.H.S.F., Lima, R.A.A., Braga-Neto, R. & Baseia, I.G. (2012) Cyathus morelensis, a rare birds nest fungus in the Brazilian Amazon
rainforest. Mycosphere 3: 880882.

http://dx.doi.org/10.5943/mycosphere/3/5/13
Cunningham, G.H. (1944) The Gasteromycetes of Australia and New Zealand. J. McInhoe, Dunedin. 236 pp.
Desvaux, N.A. (1809) Observations sur quelques genres tablir dans la famille des champignons. Journal de Botanique (Desvaux). 2:
88105.
Dissing, H. & Lange, M. (1962) Gasteromycetes of Congo. Bulletin du Jardin Botanique de lEtat 32(4): 325416.

http://dx.doi.org/10.2307/3667249
Dring, D.M. (1964) Gasteromycetes of west tropical Africa. Mycological Papers 98: 160.
Fazolino, E.P., Calonge, F.D. & Baseia, I.G. (2008) Geastrum entomophilum, a new earthstar with an unusual spore dispersal strategy.
Mycotaxon 104: 449453.
Fazolino, E. P., Trierveiler-Pereira, L., Calonge, F.D. & Baseia, I.G. (2010) First records of Clathrus (Phallaceae, Agaricomycetes) from
the Northeast Region of Brazil. Mycotaxon 113: 195202.

http://dx.doi.org/10.5248/113.195
Fischer, E. (1886) Versuch einer systematischen Uebersicht ber die bisher bekannten Phalloideen. Jahrbuch des Kniglichen Botanischen

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 251

Gartens und des Botanischen Museums zu Berlin 4: 192.

Fischer, E. (1921) Mykologische Beitrge 1820. Staheliomyces cinctus, ein neuer Typus aus der Gruppe der Phalloideen. Mitteilungen
der Naturforschung Gesellschaft in Bern 137142.
Fries, E.M. (1829) Systema mycologicum. Vol. 3. Ex Officina Berlingiana, Lund, 260 pp.
Gurgel, F.E., Silva, B.D.B. & Baseia, I.G. (2008) New records of Scleroderma from Northeastern Brazil. Mycotaxon 105: 399405.
Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic
Acids Symposium Series 41: 9598.
Hennings, P. (1904) Fungi amazonici a. cl. Ernesto Ule collecti: I. Hedwigia 43: 154186.
Hosaka, K. (2009) Phylogeography of the genus Pisolithus revisited with some additional taxa from New Caledonia and Japan. Bulletin
of the National Museum of Nature and, Series B 35(3): 151167.
Huelsenbeck, J.P. & Ronquist, F.R. (2001) MrBayes. Bayesian inference of phylogeny. Bioinformatics 17: 754755.

http://dx.doi.org/10.1093/bioinformatics/17.8.754
Junghuhn, F.W. (1840) Nova genera et species plantarum florae javanicae. Tijdschrift voor Natuurlijke Geschiedenis en Physiologie 7:
285317.
Kasuya, T., Hosaka, K., Uno, K. & Kakishima, M. (2012) Phylogenetic placement of Geastrum melanocephalum and polyphyly of
Geastrum triplex. Mycoscience 53(6): 411426

http://dx.doi.org/10.1007/S10267-012-0186-Z
Kreisel, H. (1996) A preliminary survey of the genus Phallus sensu lato. Czech Mycology 48(4): 273281.
Kreisel, H. & Dring, D.M. (1967) An emendation of the genus Morganella Zeller (Lycoperdaceae). Feddes Repertorium 74: 109122.

http://dx.doi.org/10.1002/fedr.19670740105
Kretzer, A.M. & Bruns, T.D. (1999) Use of atp6 in fungal phylogenetics: An example from the Boletales. Molecular Phylogenetics and
Evolution 13(3): 483492.

http://dx.doi.org/10.1006/mpev.1999.0680
Kppers, H. (1979) Atlas de los colores. Editorial Blume, Barcelona, 161 pp.
Lamarck, J.B.P.A.M. & De Candolle, A.P. (1805) Flore franaise, troisime dition Tome second. Chez H. Agasse, Paris, 600 pp.
Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez,
R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23: 29472948.

http://dx.doi.org/10.1093/bioinformatics/btm404
Leite, A.G., Silva, B.D.B., Arajo, R.S. & Baseia, I.G. (2007) Espcies raras de Phallales (Agaricomycetidae, Basidiomycetes) no Nordeste
do Brasil. Acta Botanica Brasilica, 21(1): 119124.

http://dx.doi.org/10.1590/S0102-33062007000100011
Leite, A.G., Assis, H.K., Silva, B.D.B., Soto, H.M.P. & Baseia, I.G. (2011) Geastrum species from the Amazon Forest, Brazil. Mycotaxon
118: 383392.

http://dx.doi.org/10.5248/118.383
Lveill, J.H. (1846) Descriptions des champignons de lherbier du Musum de Paris. Annales des Sciences Naturelles Botanique 5:
111167.
Li, T.R., Song, B. & Liu, B. (2002) Three taxa of Phallaceae in HMAS, China. Fungal Diversity 11: 123127.
Liu, B. (1984) The Gasteromycetes of China. Beihefte zur Nova Hedwigia 76: 1135.
Lloyd, C.G. (1907) The phalloids of Australasia. Mycological Writings 2: 124.
Lloyd, C.G. (1907) Mycological Notes 24. Mycological Writings 2: 293340.
Meijer, A.A.R. (2006) Preliminary checklist of the macromycetes from the state of Paran, Brazil. Boletim do Museu Botnico Municipal
(Curitiba), 68: 155.
MMA (2002) Biodiversidade Brasileira: Avaliao e identificao de reas de aes prioritrias para conservao, utilizao sustentvel,
e repartio dos benefcios da biodiversidade dos biomas brasileiros. Braslia: MMA/SBF.
Moyersoen, B. & Demoulin, V. (1996) Les Gastromyctes de Corse: taxonomie, cologie, chorologie. Lejeunia 152: 1130.
Nylander, J.A.A. (2004) MrModelTest v2. Program distributed by the editor.
Ottoni, T.B.S., Silva, B.D.B., Fazolino, E.P. & Baseia, I.G. (2010) Phallus roseus, first record from neotropics. Mycotaxon 112: 58.

http://dx.doi.org/10.5248/112.5
Patouillard, N.T. (1898) Quelques champignons de Java. Bulletin de la Socit Mycologique de France 14: 182198.
Pegler, D.N., Lsse, T. & Spooner, B.M. (1995) British puffballs, earthstars, and stinkhorns. London, Royal Botanic Gardens, Kew.
Penzig, O. (1899) Ueber Javanische phalloideen. Annales du Jardin Botanique de Buitenzorg 16: 133173.
Persoon, C.H. (1801) Synopsis methodica fungorum Pars prima. Apvd Henricvm Dieterich, Gottingen, 240 pp.
Ponce de Len, P. (1968) A revision of the Geastraceae. Fieldiana Botany 31: 303349.
Pouzar, Z. (1971) Geastrum sessile (Sow.) Pouz., the correct name for Geastrum fimbriatum Fr. (Gasteromycetes). Folia Geobotanica et

252 Phytotaxa 183 (4) 2014 Magnolia Press

CABRAL ET AL.

Phytotaxonomica 6(1): 9397.

Rick, J.E. (1906) Pilze aus Rio Grande do Sul (Brazilien). Brotria Srie Botnica 5:453.
Silva, B.D.B., Calonge, F.D. & Baseia, I.G. (2007) Studies on Tulostoma (Gasteromycetes) in the Neotropics: Some Brazilian species.
Mycotaxon 101: 4754.
Silva, B.D.B., Sousa, J.O. & Baseia, I.G. (2011) Discovery of Geastrum xerophilum from the Neotropics. Mycotaxon 118: 355359.

http://dx.doi.org/10.5248/118.355
Silva, B.D.B., Cabral, T.S., Marinho, P., Ishikawa, N.K. & Baseia, I.G. (2013) Two new species of Geastrum (Geastraceae, Basidiomycota)
found in Brazil. Nova Hedwigia 96 (34): 445456.

http://dx.doi.org/10.1127/0029-5035/2013/0089
Surez, V.L. & Wright, J.E. (1996) South American Gasteromycetes V: The genus Morganella. Mycologia 88: 655661.

http://dx.doi.org/10.2307/3761163
Swofford, D.L. (1998) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0 b. Sinauer Associates,
Sunderland, Massachusetts. 142 pp.
Tamura, K., Dudley, J., Nei, M. & Kumar, S. (2007) MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0.
Molecular Biology and Evolution 24:15961599.

http://dx.doi.org/10.1093/molbev/msm092
ter Steege, H., Pitman, N.C., Sabatier, D., Baraloto, C., Salomo, R.P., Guevara, J.E., Phillips, O.L., Castilho, C.V., Magnusson, W.E.,
Molino, J.F., Monteagudo, A., Nez Vargas, P., Montero, J.C., Feldpausch, T.R., Coronado, E.N., Killeen, T.J., Mostacedo, B.,
Vasquez, R., Assis, R.L., Terborgh, J., Wittmann, F., Andrade, A., Laurance, W.F., Laurance, S.G., Marimon, B.S., Marimon, B.H.
Jr., Guimares Vieira, I.C., Amaral, I.L., Brienen, R., Castellanos, H., Crdenas Lpez, D., Duivenvoorden, J.F., Mogolln, H.F.,
Matos, F.D., Dvila, N., Garca-Villacorta, R., Stevenson Diaz, P.R., Costa, F., Emilio, T., Levis, C., Schietti, J., Souza, P., Alonso, A.,
Dallmeier, F., Montoya, A.J., Fernandez Piedade, M.T., Araujo-Murakami, A., Arroyo, L., Gribel, R., Fine, P.V., Peres, C.A., Toledo,
M., Aymard, C.G.A., Baker, T.R., Cern, C., Engel, J., Henkel, T.W., Maas, P., Petronelli, P., Stropp, J., Zartman, C.E., Daly, D.,
Neill, D., Silveira, M., Paredes, M.R., Chave, J., Lima Filho Dde, A., Jrgensen, P.M., Fuentes, A., Schngart, J., Cornejo Valverde,
F., Di Fiore, A., Jimenez, E.M., Peuela Mora, M.C., Phillips, J.F., Rivas, G., van Andel, T.R., von Hildebrand, P., Hoffman, B.,
Zent, E.L., Malhi, Y., Prieto, A., Rudas, A., Ruschell, A.R., Silva, N., Vos, V., Zent, S., Oliveira, A.A., Schutz, A.C., Gonzales,
T., Trindade Nascimento, M., Ramirez-Angulo, H., Sierra, R., Tirado, M., Umaa Medina, M.N., van der Heijden, G., Vela, C.I.,
Vilanova Torre, E., Vriesendorp, C., Wang, O., Young, K.R., Baider, C., Balslev, H., Ferreira, C., Mesones, I., Torres-Lezama, A.,
Urrego Giraldo, L.E., Zagt, R., Alexiades, M.N., Hernandez, L., Huamantupa-Chuquimaco, I., Milliken, W., Palacios Cuenca, W.,
Pauletto, D., Valderrama Sandoval, E., Valenzuela Gamarra, L., Dexter, K.G., Feeley, K., Lopez-Gonzalez, G. & Silman, M.R.
(2013) Hyperdominance in the Amazonian Tree Flora. Science 342: 1243092-11243092-9.
Trierveiler-Pereira, L., Gomes-Silva, A.C. & Baseia, I.G. (2009) Notes on gasteroid fungi in the Brazilian Amazon rainforest. Mycotaxon
110: 7380.

http://dx.doi.org/10.5248/110.73
Trierveiler-Pereira, L., Gomes-Silva, A.C. & Baseia, I.G. (2011) Observations on gasteroid Agaricomycetes from the Brazilian Amazon
rainforest. Mycotaxon 118:273282.

http://dx.doi.org/10.5248/118.273
Vigalys, R. & Hester, M. (1990) Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several
Cryptococcus species. The Journal of Bacteriology 172(8): 42384246.
Veloso, H.P., Rangel-Filho, A.L.R. & Lima, J.C.A. (1991) Classificao da vegetao brasileira, adaptada a um sistema universal. Instituto
Brasileiro de Geografia e Estatstica, Rio de Janeiro, BR.
Vittadini, C. (1842) Monographia Lycoperdineorum. Memorie della Reale accademia delle scienze di Torino 5: 145238.
Zeller, S.M. (1948) Notes on certain Gasteromycetes, including two new orders. Mycologia 40(6): 639668.

http://dx.doi.org/10.2307/3755316

A new species gasteroid fungi (Basidiomycota)

Phytotaxa 183 (4) 2014 Magnolia Press 253