![FIGURE . Phellinotus species. P. neoaridus. (A) holotype on specific host (lenticels of Caesalpinia sp. in detail); (B) applanate to triquetrous basidioma with a distinct ochre-orange margin (URM80410); (C) ungulate with strongly rimose upper surface and brownish margin (URM80641) (Bars = 5 cm); (D): isotype context and tubes lacking tissue layer between the tubes, in detail (D*) the black line near the upper surface (Bars = 1 cm); (E–G) basidiospores, white arrows are indicating the adaxially flattened surface [(E) lactophenol, (F) KOH+, (G) CB±; Bars = 5 µm]; P. piptadeniae. (H) paratype (IAC4365); (I) context in detail showing the distinct black line in the paratype; (J) mature triquetrous, developing applanate basidiomata on Piptadenia spp. (URM 80768) (Bars = 5 cm); (L) black line and tissue layer between the tubes are showing in detail in L and L* (Bars = 1 cm); (M–O) basidiospores, white arrows are indicating the adaxially flattened surface [(M): lactophenol, (N) KOH+, (O) CB±; Bars = 10 µm]. Photos of basidiomata (A, B, C, D, H, I, J) by Elisandro](https://www.researchgate.net/profile/Tatiana-Gibertoni/publication/301692762/figure/fig2/AS:373694636544002@1466107152961/FIGURE-Phellinotus-species-P-neoaridus-A-holotype-on-specific-host-lenticels-of_Q320.jpg)
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Phytotaxa 000 (0): 000–000
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Genevieve Gates: 29 Apr. 2016; published: xx xxx. 2016
http://dx.doi.org/10.11646/phytotaxa.00.0.0
1
Phellinotus, a new neotropical genus in the Hymenochaetaceae (Basidiomycota,
Hymenochaetales)
ELISANDRO RICARDO DRECHSLER-SANTOS1,*, GERARDO LUCIO ROBLEDO2,3*, NELSON C. LIMA-
JÚNIOR4, ELAINE MALOSSO4, MATEUS A. RECK1, TATIANA B. GIBERTONI4, MARIA A. DE QUEIROZ
CAVALCANTI4 & MARIO RAJCHENBERG5
1Laboratório de Micologia, Departamento de Botânica, Universidade Federal de Santa Catarina, CEP: 88040-900 Florianópolis, SC,
Brazil;
2Laboratorio de Micología, Instituto Multidisciplinario de Biología Vegetal, CONICET - Universidad Nacional de Córdoba, CC 495,
CP 5000 Córdoba, Argentina;
3FungiCosmos, Av. General Paz 154, 4º piso, oficina 4, Cordoba, Argentina;
4Departamento de Micologia, Universidade Federal de Pernambuco, CEP: 50760-420 Recife, PE, Brazil;
5Protección Forestal, Centro Forestal CIEFAP, C.C. 14, 9200 Esquel, Chubut, Argentina.
* Both authors contributed equally to the research and should be considered co-first authors
Correspondence to: Elisandro R. Drechsler-Santos. E-mail: e.ricardo@ufsc.br; drechslersantos@yahoo.com.br)
Abstract
A new poroid genus with two conspicuous and common species growing on living Fabaceae trees is described from the
Seasonally Dry Tropical Forests biome of Brazil. Both taxa in this forest pathogen genus resemble Phellinus rimosus macro-
scopically, but are distinguished by a dimitic hyphal system with skeletal hyphae present only in the trama of the tube layer
while the context remains monomitic, and by the ellipsoid, thick-walled, adaxially flattened, yellow basidiospores that turn
chestnut brown in KOH solution. Molecular and morphological studies of Brazilian specimens macroscopically similar to
the Phellinus rimosus species complex were carried out to solve their phylogenetic relationships among the Hymenochae-
taceae. Phellinotus gen. nov. with P. neoaridus sp. nov. as the genus type and P. piptadeniae comb. nov. are presented and
described. Phylogenetically, Phellinotus is closely related to Arambarria, Inocutis, Fomitiporella and other taxonomically
unresolved terminal clades, and unrelated to Fulvifomes and Phylloporia. Phellinotus and other genera of poroid Hymeno-
chaetaceae that lack setae or setal hyphae and produce thick-walled, colored (pale yellow to rusty brown) basidiospores form
a phylogenetic group here named the ‘phellinotus clade’. Our results indicate the need to include taxa from unexplored areas
in order to get a thorough understanding of the phylogeny of the Hymenochaetaceae.
Key words: forest pathogen, Fulvifomes, molecular phylogeny, mycodiversity, Phellinus rimosus complex, polypores, tax-
onomy
Introduction
Hymenochaetaceae Donk (Hymenochaetales Oberw., Agaricomycetes Doweld) comprises about 400 species and, as
originally circumscribed (Donk 1964), is not a monophyletic family (Larsson et al. 2006). Traditionally, the family has
been characterized by wood-decaying fungi with styrylpyrone contents that are responsible for a positive xanthochroic
reaction on the basidiomata (e.g., a black coloration of the surface and tissues in 5% KOH solution), simple-septate
hyphae and setal elements (Cannon & Kirk 2007). Most poroid Hymenochaetaceae were placed in Phellinus Quél.
(1886:172) and Inonotus P. Karst. (1879:39), except for species of Coltricia Gray (1821:644), Phylloporia Murrill
(1904:141) and Cyclomyces Kunze ex Fr. (1830:512), which have macro- and micro-morphological characters that
allow for their ready identification (Ryvarden 1991). Species of Phellinus and Inonotus are defined primarily by the
type of hyphal system, either monomitic or dimitic (with skeletal hyphae) hyphal systems, and the annual or perennial
habit of the basidiomata (Pilát 1936–42, Cunningham 1947). Species with a dimitic hyphal system and perennial
basidioma were placed in Phellinus whereas species with a monomitic hyphal system and annual basidioma were
identified as Inonotus (Fiasson & Niemelä 1984, Corner 1991, Wagner & Fischer 2001). In this context, several
smaller genera proposed by Murrill at the beggining of the 20th century were considered as synonyms of Phellinus and
DRECHSLER-SANTOS ET AL.
2 • Phytotaxa 000 (0) © 2016 Magnolia Press
Inonotus (Pilát 1936–42, Cunningham 1947). Among the perennial species (Phellinus s.l.) those with pileate basidioma,
Pyropolyporus Murrill (1903:109) and Porodaedalea Murrill (1905:367) were proposed on account of their poroid
and daedaloid hymenophores, respectively and those with resupinate basidioma, Fomitiporia Murrill (1907:7) and
Fomitiporella Murrill (1907:12) were proposed on account of their hyaline and brown spores, respectively. Fulvifomes
Murrill (1914:49) was proposed to accommodate Pyropolyporus species with brown spores. Among the annual species
(Inonotus s.l.), Fuscoporia Murrill (1907:3) and Fuscoporella Murrill (1907:6) were distinguished by their hyaline and
brown spores, respectively (Murrill 1903, 1904, 1905, 1907, 1914).
Nevertheless, before the advent of molecular phylogenetics, authors neglected those names and accepted Phellinus
and Inonotus in a wide sense (Lowe 1957, Reid 1963, Pegler 1964, Gilbertson & Ryvarden 1986 and 1987, Wright &
Blumenfeld 1984, Larsen & Cobb-Poulle 1990, Ryvarden 2004, 2005). However, Phellinus and Inonotus were shown
to be polyphyletic by molecular phylogenetics and reinforced the observation that the type of hyphal system and
basidioma persistence are not valid generic traits in the Hymenochaetaceae (Wagner & Fischer 2002).
A revision of the European taxa of Phellinus and Inonotus was proposed by Fiasson & Niemelä (1984). They
identified smaller species groups that they recognized as morphologically distinct genera, including Inocutis Fiasson &
Niemelä (1984:24), a new genus based on morphological, cultural, chemical, and nuclear behavioural data. Since then,
Fiasson & Niemelä’s (1984) results were corroborated by Niemelä et al. (2001) and Wagner & Fischer (2001, 2002)
through molecular and morphological methods. These studies and others (e.g. Zhou & Qin 2013) supported the genera
Aurificaria D.A. Reid (1963:278), Fomitiporella, Fomitiporia, Fuscoporia, Inocutis, Inonotus s.s., Mensularia Lázaro
Ibiza (1916:736), Phellinidium (Kotl.) Fiasson & Niemelä (1984:25), Phellinopsis Y.C. Dai (2010:309), Phellinus s.s.,
Phellopilus Niemelä T., Wagner & M. Fischer (2001:53), Phylloporia Murrill, Porodaedalea and Pseudoinonotus T.
Wagner & M. Fischer (2001:781).
Larsson et al. (2006) presented a molecular phylogenetic analysis of the Hymenochaetales which supported several
different genera of poroid Hymenochaetaceae. Other studies have contributed to the phylogenetic relationships among
the Hymenochaetaceae, such as those by Decock et al. (2005, 2006, 2007), Jeong et al. (2005), Dai et al. (2008), Dai
(2010), Tura et al. (2012), Tian et al. (2013), Zhou & Qin (2013), Parmasto et al. (2014), Zhou (2014a), Rajchenberg
et al. (2015), Zhou (2015) and Zhou et al. (2015). However, the classification of this group is incomplete because, as
with other regions, neotropical taxa were often absent in the phylogenetic studies.
Research on poroid Hymenochaetaceae in the Caatinga dry woodlands, a northeastern Brazilian part of the
Seasonally Dry Tropical Forests biome - SDTFs (Särkinen et al. 2011), recovered several specimens that macroscopically
resemble the Phellinus rimosus species complex (Ryvarden 2004). Specimens were identified as Phellinus piptadeniae
Teixeira (1950:118), repeatedly recorded on Piptadenia Benth. (1840:135), and P. rimosus (Berk.) Pilát (1940:80)
s.l. on Caesalpinia L. (1753:380) (Drechsler-Santos et al. 2010). Species in the P. rimosus species complex are
characterized by having woody, pileate basidiomata with a cracked (rimose) pileus, a dimitic hyphal system throughout
the basidioma, a lack of setae, and ellipsoid, thick-walled, chestnut to rusty brown basidiospores (Fiasson & Niemelä
1984). These morphological features indicate that P. rimosus and similar species belong in Fulvifomes, as proposed
by Fiasson & Niemelä (1984) and confirmed by Dai (2010). The aim of this study was to investigate the identity of P.
rimosus s.l. and P. piptadeniae from Caatinga, Brazil, and to investigate the phylogenetic relationship of these species
to Fulvifomes and other Hymenochaetaceae.
Materials and methods
Collections
Collecting was carried out in the Caatinga dry woodlands in the states of Alagoas, Ceará, Paraíba, Pernambuco, Rio
Grande do Norte and Sergipe (Brazil) and specimens were deposited in the herbaria at CORD, FLOR, HTSA, O,
URM, and Centro Forestal CIEFAP (Esquel, Argentina). Herbarium codes follow Thiers (2015).
Morphological studies
Basidiomata were macro- and micro-morphologically analyzed. Microscopic observations were carried out on
basidioma sections stained with 3–5% KOH and 1% phloxine, neutral lactophenol, cotton blue, or Melzer’s reagent
(Largent et al. 1977, Ryvarden 1991). Measurements were taken from 3% KOH preparations, following a sampling
of 40 (n = 40) structures such as pores/mm, basidiospores, and diameter of hyphae. The size of microscopic elements
is given as values (or an interval) followed by 5% variation in parentheses. Abbreviation and symbols are KOH+ =
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 3
changing color (reacting) in KOH; IKI- = not reacting in Melzer’s reagent; CB± = weakly cyanophilous; (!) = type
studied; (≡) = homotypic synonym; (=) = heterotypic synonym; and Q = quotient of basidiospores length (L) / width
(W), L = mean of length (arithmetical mean of all basidiospores), and W = mean of width; ave. = arithmetic average;
ave.Q = Q arithmetic average. To access and determine the hyphal system under microscopy, sections from the trama
of tube layer and context of basidiomata under the pilear surface and between the tube layers were carefully dissected
under a stereomicroscope after incubation in 3‒5% NaOH for 48h at 40°C (Decock et al. 2010). Reference material
of Phellinus piptadeniae [BRAZIL. São Paulo: Campinas, Bosque dos Jequitibás, on living Piptadenia communis
Beth. (1841:337–338), 12 September 1943, A.R. Teixeira and P.R. Santos sine numerus (sn) (paratype IAC 4365)],
Pyropolyporus robiniae Murrill (1903:114) [USA. Ohio: January 1957, Lloyd 223 (lectotype NY 743007), Phylloporia
parasitica Murrill (1904:141) [COLOMBIA. Santa Marta: near Bonda, 16 December 1898, C. F. Baker sn (lectotype K-
M 00705227)], Arambarria destruens Rajchenb. & Pildain (2015:759) [ARGENTINA. Chubut: Lago Puelo National
Park, W arm of lago Puelo, oriental slope of Valle de las Lágrimas, Los Tineos stream, 10 May 1996, M. Rajchenberg
11172 (holotype BAFC 34575)], and Fomitiporella umbrinella (Bres.) Murrill (1907:13) [BRAZIL. Santa Catarina:
Blumenau, Parque Nacional da Serra do Itajaí, 27º02’20” S, 49º03’24” W, 12 June 2012, MAB Silva & al. 271 (FLOR
49263)] were used for morphological comparisons.
DNA extraction
Fragments of basidioma (30 mg) were ground with a pestle in a porcelain mortar containing liquid nitrogen. The
resulting powder was transferred to a 1.5 mL pre-warmed (65ºC) microcentrifuge tube containing 700 μL of extraction
buffer [CTAB 2%, 100 mM Tris-HCl pH 8, 1.4 M NaCl, 20 mM EDTA, 1% PVP (Rogers & Bendich 1985)] and
incubated at 65ºC for at least 30 minutes. DNA was extracted once with chloroform-isoamyl alcohol (24:1), precipitated
with isopropanol, washed with 70% ethanol, and re-suspended in 50 μL ultrapure water (Góes-Neto et al. 2005).
Polymerase chain reaction and sequencing
Reaction mix and parameters for PCR amplification of the full ITS (Internal transcribed spacer of nuclear ribosomal
RNA gene) regions followed Kaliyaperumal & Kalaichelvan (2008) using the primers ITS1 and ITS4 (White et al.
1990). For the nLSU (Large sub-unit of nuclear ribosomal RNA gene), amplification was carried out with parameters
and reagent concentrations given by Góes-Neto et al. (2005) using the primers LR0R and LR7 (Moncalvo et al. 2000).
The amplicons were purified using the PureLink PCR Purification Kit (Invitrogen) and the purified products were
sequenced at the Human Genome Research Center of the Universidade de São Paulo (USP, Brazil) in an ABI-310
Capillary Sequencer (PerkinElmer, Wellesley Massachusetts, USA). Primers LR0R/LR5 and ITS1/ITS4 were used for
sequencing the nLSU and ITS, respectively. The sequences were deposited in the GenBank (Benson et al. 2009).
Phylogenetic analysis
Seven ITS and seven nLSU sequences obtained in this study were used to search for sequences of the closest related
taxa, using BLASTn searches in the GenBank (see Appendix 1). Most of the sequences and species in this study
represent the currently accepted genera of Hymenochaetaceae based on phylogenetic concepts (Wagner & Fischer 2002,
Larsson et al. 2006, Rajchenberg et al. 2015). ITS and nLSU sequences obtained in this study and those retrieved from
GenBank are added to the final matrix. Trichaptum sector (Ehrenb.) Kreisel (1971:84) was defined as the outgroup,
based on previous papers that showed this genus as related to Hymenochaetaceae within Hymenochaetales (Larsson et
al. 2006). All specimens and sequences used in this study are listed in the Appendix 1.
The final ITS and nLSU datasets were then automatically aligned with Mafft v.7 (Katoh & Standley 2013) under
the E-INS-I and G-INS-I strategies, respectively. The fisrt strategy is suitable for regions with multiple conserved
domains and long gaps, while the second is for regions with global homology. Both of the alignments were manually
adjusted as necessary with MEGA v.6.0 (Tamura et al. 2013). Unreliably aligned positions in the ITS alignment were
detected using GBLOCKS v.0.91b (Castresana 2000), using the following parameters: minimum number of sequences
for a conserved position and for a flank position, both set to 43; maximum number of contiguous nonconserved
positions set to 6; minimum length of a block set to 3; and allowed gap positions set to with half. The final alignment
was then divided in four partitions: ITS 1, 5.8S, ITS 2 and nLSU. The best nucleotide substitutions model for each
partition was estimated using the AIC (Akaike Information Criterion), as implemented in jModelTest2 v.2.1.6 (Darriba
et al. 2012, Guindon & Gascuel 2003).
Two phylogenetic routines were used to reconstruct the relationships hypothesis, using the concatenated dataset
with the four partitions: Maximum Likelihood (ML) and Bayesian Inference (BI). ML analysis was carried out as
implemented in RAxML v.8.1.24 (Stamatakis 2014), available in the CIPRES science gateway (Miller et al. 2010).
DRECHSLER-SANTOS ET AL.
4 • Phytotaxa 000 (0) © 2016 Magnolia Press
The analysis first involved 100 ML hill-climbing searches, each one starting from one randomized stepwise addition
parsimony tree, under a GTRGAMMAI model, with all parameters estimated by the software. Only the best-scored
likelihood tree from all searches was kept. To access the reliability of the nodes, multiparametric bootstrapping
replicates under the same model were computed, allowing the program to halt bootstrapping automatically by the
autoMRE option. The bootstrap values were then annotated to the best likelihood tree find. An additional file with the
partitions was informed to force the software to estimate the parameters for each partition independently, through all
steps of the analyses.
To check for incongruences among the partitions, each partition was analyzed under ML separated under the same
parameters described above, comparing the resulting topologies and looking for conflicts only involving branches with
BS ≥ 70% (Reeb et al. 2004). A conflict was assumed to be significant if two different relationships for the same set of
taxa (one being monophyletic and the other non-monophyletic) were observed in rival trees.
BI was performed in the program Mr. Bayes v.3.2.6 (Ronquist et al. 2012). The analysis was implemented by two
MCMC independent runs, each one starting from random trees and with four simultaneous independent chains, performing
15,000,000 generations, keeping one tree every 1000th generation. Four rate categories were used to approximate the
gamma distribution, and the nucleotide substitutions rates were fixed to the estimated values. The first 20% of the sampled
trees was discarded as burn-in, checked by the convergence criterion (frequencies of average standard deviation of split
<0.01) in Tracer v.1.6 (Rambaut et al. 2014), while the remaining ones were used to reconstruct a 50% majority-rule
consensus tree and to estimate Bayesian Posterior Probabilities (BPP) of the branches.
A node was considered to be strongly supported if it showed a BPP ≥ 0.95 and/or BS ≥ 90%, while moderate
support was considered when BPP ≥ 0.9 and/or BS ≥ 70%. The alignment was deposited in TreeBASE (http://www.
treebase.org/treebase/index.html), under accession ID: 18406.
Results
Molecular analyses
The aligned matrix of the ITS 1 dataset was 160 bp, for the 5.8s was 155 bp, for the ITS 2 was 195 bp, while for nLSU
dataset it was 853 bp, resulting in a final gapped matrix of 1363 bp.
The concatenated dataset included sequences from 82 fungal specimens representing the main genera of
Hymenochaetaceae, and one representing the outgroup. According to the preceding analyses, GTR +I+G (ITS1), JC
(5.8s), K80+G (ITS2) and GTR+I+G (nLSU) were chosen as the best-fit substitution models for each dataset. During
the BI analyses, the indepentent Markov chains reach stability after 921,000 generations, which confirmed that the used
20% burn-in value was appropriate. The bootstopping criteria of the ML algorithm found that 252 pseudoreplicates
were sufficient for the dataset.
The phylogenetic reconstructions based on ML and BI analyses for the combined ITS+nLSU datasets did not
show major conflicts in the tree topology and were mostly congruent, which allowed us to combine them (Fig. 1).
Overall, most of the current genera were recognized in the analyses. The studied specimens from Caatinga, previously
determined as P. rimosus s.l. and P. piptadeniae (Drechsler-Santos et al. 2010), clustered in the same generic clade with
high support (BS=99, BPP=1), however separated from the currently known genera. This clade is here recognized as a
new genus Phellinotus. It is nested in a more inclusive clade composed of poroid members of the Hymenochaetaceae
in which setal elements are absent and possess colored and thick-walled basidiospores, e.g., Arambarria Rajchenb.
& Pildain (2015: 759), Fomitiporella, Fulvifomes, Inocutis, and Phylloporia, among other taxonomically unresolved
lineages. This more inclusive clade, with significant support in both analyses, moderate in ML (BS= 87) and strong
in BI (BPP=1), is here named the ‘phellinotus clade’. Despite some lower support values in some basal branches, the
overall topology from both ML and BI analyses is quite similar within this clade. Phellinotus appears to be the sister
group of specimens identified as Inocutis dryophila (Berk.) Fiasson & Niemelä (1984:25), but this relationship was
not significantly supported by the data analyzed. The relationship with other genera of the ‘phellinotus clade’ remains
unresolved, but species of Arambarria and Fomitiporella and specimens determined as Fulvifomes chinensis (Pilát)
Y.C. Dai, F. inermis (Ellis & Everh.) Y.C. Dai (2010:187), Inonotus tenuissimus H.Y. Yu, C.L. Zhao & Y.C. Dai
(2013:64), and unidentified isolates CIEFAPcc88 and 107 (Rajchenberg et al. 2015) appear to be closely related to
Phellinotus (Fig. 1). Also, the genera Fulvifomes s.s. and Phylloporia were recovered within the ‘phellinotus clade’.
The data also showed that the two species under study are closely related but can be considered as two distinct species.
Phellinus rimosus s.l. specimens from Caatinga, Brazil were unrelated to Fulvifomes s.s. (P. rimosus). Therefore, they
were assigned to a new species Phellinotus neoaridus.
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 5
FIGURE 1. Phylogram of Hymenochaetaceae recovered from ITS (ITS1, 5.8S, ITS2) and nucLSU regions, inferred by Maximum
likelihood analysis (log likelihood -16996.695988). Support values along branches are bootstrap from ML (≥ 50, before slash) and
Bayesian posterior probability (≥0.7, after slash). Trichaptum sector was used to root the tree.
Taxonomy
Phellinotus Drechsler-Santos, Robledo & Rajchenb. gen. nov.
Typus generis:—Phellinotus neoaridus Drechsler-Santos & Robledo.
Mycobank no:—MB804717
Etymology:—“Phelli” from Phellinus + “notus” (ear) from Inonotus, in reference to Phellinus s.l. and Inonotus s.l., which traditionally
were considered dimitic and monotic, respectively. Phellinotus combines both dimitic and monomitic hyphal systems in different
parts of basidiomata.
Diagnosis:—Basidioma pileate, annual to perennial; hymenophore poroid. Hyphal system dimitic with skeletal hyphae found only in
trama of tube layer; generative hyphae with simple septa, skeletal hyphae aseptate. Basidiospores ellipsoid, adaxially flattened, thick-
walled, pale yellow, becoming chestnut brown in KOH solution. Setae and cystidioles absent.
DRECHSLER-SANTOS ET AL.
6 • Phytotaxa 000 (0) © 2016 Magnolia Press
Description:—Basidioma annual to perennial, pileate, applanate to ungulate, fulvous brown to dark brown. Pileus
brown to blackened, rugose to rimose. Context with a black line near/below the upper surface, distinct or indistinct.
Tubes stratified, with or without contextual tissue layer between them. Hymenophore poroid, pores irregularly rounded,
fulvous brown to deep brown. Hyphal system dimitic with skeletal hyphae restricted to the trama of tube layer;
contextual generative hyphae in different stages of development, thin- to thick-walled, first regularly septate, branched,
becoming sclerified and some portions of thick-walled hyphae sparsely simple-septate; tramal hyphae with simple-
septate generative and skeletal hyphae. Setae and other sterile elements absent. Basidia not observed. Basidiospores
broadly ellipsoid to ellipsoid, adaxially flattened, smooth, thick-walled, yellow in lactophenol, becoming chestnut
brown in KOH solution, weakly cyanophilous, IKI-.
Habitat and distribution:—Found on living members of the family Fabaceae, with host specialization (Teixeira
1950, Drechsler-Santos et al. 2010, as Phellinus rimosus and P. piptadeniae). So far known from the South American
Seasonally Dry Tropical Forest (SDTF) biome, from the seasonally dry forests of the Atlantic Forest (states of Santa
Catarina and São Paulo) and Caatinga (states of Alagoas, Bahia, Ceará, Paraíba, Pernambuco, Rio Grande do Norte
and Sergipe) domains of Brazil, and from lowland SDTF of northwestern Peru (Drechsler-Santos et al. 2010, 2013,
Salvador-Montoya et al. 2015).
TABLE 1. Morphological features of poroid Hymenochaetaceae genera.
Genera Substrate Basidioma Hyphal
System
Setae Basidiospores
Shape Tube
layers
Shape Colour Wall/Dextrinoid/
KOH reaction
Arambarria DW RES/EFR AN MO Lacking EBE; FLT Yellow Thick / - / +
Asterodon CW RES/EFR AN DI Hymenial;
simple/branched
EBE Hyaline Thin / - / -
Aurificaria DW PIL AN MO Lacking EBE; FLT Yellow Thick / - / +
Cylindrosporus DW PIL AN MO Hymenial CYL Hyaline Thin / - / -
Fomitiporella DW RES/EFR P MO/DI Lacking EBE; FLT Brown Thick / - / +
Fomitiporia DW/CW RES/EFR/
PIL
P MO/DI Lacking or rare;
hymenial
GLB Hyaline Thick / + / -
Fulvifomes DW RES/EFR/
PIL
P DI Lacking GLB/
EBE; FLT
Yellow Thick / - / +
Fuscoporia DW/CW RES/EFR/
PIL
AN/P MO/DI Hymenial EBE/ALL/
CYL
Hyaline Thin / - / -
Inocutis DW PIL AN MO Lacking EBE; FLT Yellow/
brown
Thick / - / -
Inonotopsis CW RES AN MO Lacking EBE Hyaline Thin / - / -
Inonotus s.s.DW RES/EFR/
PIL
AN/P MO/DI Mostly presente;
hymenial/Setal
hyphae
EBE/GLB Yellow/
brown
Thin-thick / - / -
Mensularia DW RES/EFR/
PIL
AN MO Hymenial/
tramal
EBE Hyaline Thick / + / -
Nothophellinus DW PIL P DI-TRI Lacking CYL/OBC Hyaline Thin / - / -
Onnia CW SST AN MO Tramal to
hymenial
EBE/GLB Hyaline Thin / - / -
Phellinidium CW RES/EFR/
PIL
AN/P MO Setal hyphae EBE/ALL/
CYL
Hyaline Thin / - / -
...Continued on next page
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 7
TABLE 1. (Continued)
Genera Substrate Basidioma Hyphal
System
Setae Basidiospores
Shape Tube
layers
Shape Colour Wall/Dextrinoid/
KOH reaction
Phellinus s.s.DW RES/EFR/
PIL
P DI Hymenial EBE/GLB Hyaline Thin-thick / - / -
Phellinopsis DW RES/EFR/
PIL
P DI Arising from
trama
EBE Hyaline/
Yellow
Thick / - / -
Phellinotus DW PIL P DI Lacking EBE; FLT Yellow Thick / - / +
Phellopilus CW RES/EFR/
PIL
P DI-TRI Hymenial OBC/CYL Hyaline Thin / - / -
Phylloporia DW RES/EFR/
PIL
AN/P MO/DI Lacking EBE/
CYL; FLT
Pale
yellow
Thick / - / -
Porodaedalea CW RES/EFR/
PIL
P DI Hymenial EBE/GLB Hyaline Thin-thick / - / -
Pseudoinonotus DW/CW PIL AN MO/DI Hymenial GLB Hyaline Thick / + / -
Pyrrhoderma DW PIL/SST AN/P MO Lacking GLB Hyaline Thin / - / -
Sanghuangporus DW EFR/PIL P DI Hymenial EBE/GLB Pale
yellow
Thick / - / -
Tropicoporus DW RES/EFR/
PIL
AN/P DI Hymenial EBE/GLB Pale
yellow
Thick / - / -
DW = deciduous wood, CW = coniferous wood; DI = dimitic, MO = monomitic, TRI = trimitic, P = perennial, AN = annual-biannual,
RES = resupinate, EFR = effused-reflexed, PIL = pileate, SST = substipitate to stipitate, EBE = ellipsoid-broadly ellipsoid, FLT =
flattened, GLB = globose-subglobose, ellipsoid, ALL = allantoid, CYL = cylindric-subcylindric, OBC = obclavate, (+) or (-) = positive
or negative reaction in the basidiospores wall. Genera of the ‘phellinotus clade’ are presented in bold type.
Remarks:—Phellinotus is characterized by a pileate basidioma often with a rimose pileus, a dimitic hyphal system
with skeletal hyphae only present in the trama of the tube layer, and adaxially flattened, ellipsoid, thick-walled, pale
yellow basidiospores that turn chestnut brown in KOH solution. The absence of setae and setal hyphae and the presence
of colored (pale yellow to brown) and thick-walled basidiospores indicate that Phellinotus is morphologically similar
to Arambarria, Aurificaria, Fomitiporella, Fulvifomes, Inocutis, and Phylloporia (Larsson et al. 2006, Rajchenberg
et al. 2015). Fomitiporella and Fulvifomes have a dimitic hyphal system throughout the basidioma, while species of
Arambarria, Aurificaria and Inocutis are monomitic. Additionally, some species of Fulvifomes have mainly globose
to subglobose, yellowish basidiospores (Wagner & Fischer 2002), that become ferruginous to fulvous in KOH (5.5‒6
× 4.5‒5.5 µm, as observed in the lectotype of Pyropolyporus robiniae Murrill), while Phylloporia produces smaller,
light yellow basidiospores (Ryvarden 2004), as observed in the lectotype of P. parasitica Murrill. Fomitiporella also
produces basidiospores that turn darker in KOH, as verified in reference material of F. umbrinella, the genus type,
collected in the type locality (FLOR49263); however, species in this genus develop resupinate basidiomata and are
saprobes (Murrill 1907, Teixeira 1994, Zhou 2014b) (Table 1). This morphologically and molecularly related (Fig.
1) group of genera, e.g., Arambarria, Fomitiporella, Fulvifomes, Inocutis, Phylloporia, and other taxonomically
unresolved lineages, is named here as the ‘phellinotus clade’. Inonotopsis Parmasto (1973:12) and Pyrrhoderma
Imazeki (1966:4) are poroid members of the Hymenochaetaceae that also lack setae and setal hyphae; however, they
are phylogenetically unrelated and morphologically differ from members of the ‘phellinotus clade’ by producing thin-
walled basidiospores.
Phellinotus neoaridus Drechsler-Santos & Robledo sp. nov. (Figs 2A–G, 3A, C, E, Table 2)
Typus:—BRAZIL. Pernambuco: Serra Talhada, Estação Experimental do IPA, on living caatingueira tree (Caesalpinia sp.), 7º53’29” S,
38º18’17” W‒490m alt., 09 December 2008, Drechsler-Santos DS105PE (holotype URM 80362, isotype FLOR 53152).
DRECHSLER-SANTOS ET AL.
8 • Phytotaxa 000 (0) © 2016 Magnolia Press
Mycobank no:—MB805901
Etymology:—neoaridus, in reference to the neotropical semiarid region of Brazil covered by Caatinga, a seasonally dry forest of the
SDTFs biome (Särkinen et al. 2011), where the species was first encountered.
Diagnosis:—Basidioma pileate, annual to perennial, distinctly rimose; black lines near the upper surface of the pileus present; tubes
distinctly stratified, hymenophore poroid (3‒6/mm). Hyphal system dimitic with skeletal hyphae restricted to trama of tube layer.
Basidiospores ellipsoid (5‒7 × 4‒5.5 µm), adaxially flattened, thick-walled, yellow, chestnut brown in KOH. Pathogenic on
Caesalpinia spp.
Description:—Basidioma mostly perennial, more rarely annual to bi-annual, pileate, projecting up to 150 mm, 100
mm wide and 80 mm thick at base, first applanate, triquetrous to strongly ungulate with age, sometimes with a basal
umbo; hard; usually solitary or in groups of several basidiomata widespread along the substrate; margin applanate to
convex, always distinct from older parts, cream to ochre orange at first, then turning brown with age. Pileus variable
in color, first cream to reddish yellow or olive to black, dull when rimose, azonate then sulcate, and finally strongly
rimose. Context 11‒21 mm thick in young, 3‒5 mm thick in ungulate specimens, weakly zonate near the base to
azonate towards the margin, with black lines near the upper surface of the pilei, granular core variably present, pale to
dark brown, becoming black with KOH, a thick gray to black crust present above contex. Tubes stratified, dark reddish
brown, without distinct tissue development between layers.
TABLE 2. Detailed morphology (n=40) of Phellinotus species and their hosts.
Phellinotus
species/specimens
Basidioma Context
dark line
Pore surface Basidiospores Skeletal
hypha
(µm long)
Hosts
Pores/mm ave. size (µm) Shape
Phellinotus neoaridus P AU Indistinct 3‒6 5.6‒6 × 4.2‒5BE-E 110‒560 on Caesalpinia
spp.
URM 80494 P AU Indistinct 3‒4 (‒5) 5.6 × 4.2 BE-E 150‒300 C. pyramidalis
Robledo 1974 P AU Indistinct 3‒4 (‒5) 5.7 × 4.4 BE-E 110‒250 Caesalpinia sp.
URM 80299 P U Indistinct (3‒) 4‒6 5.9 × 4.5 BE-E 205‒554 Caesalpinia sp.
URM 80411 P AU Indistinct (4‒) 5‒6 5.8 × 4.8 BE-E 237‒560 Caesalpinia sp.
URM 80419 P U Indistinct 4‒6 5.7 × 4.5 BE-E 189‒339 Caesalpinia sp.
URM 80422 P AU Indistinct 4‒5 (‒6) 5.9 × 4.9 BE-E 209‒489 Caesalpinia sp.
URM 80536 P T Indistinct (4‒) 5‒6 5.9 × 4.9 BE-E 220‒524 Caesalpinia sp.
URM 80577 P U Indistinct 4‒5 (‒6) 5.6 × 4.6 BE 175‒372 Caesalpinia sp.
URM 80579 P AU Indistinct (4‒) 5‒6 6.0 × 5.0 BE-E 264‒446 Caesalpinia sp.
URM 80769 P U Indistinct (4‒) 5‒6 6.0 × 4.8 BE 245‒426 Caesalpinia sp.
URM 80362 (holotype) P U Indistinct (3‒) 4‒5 (‒6) 5.9 × 4.3 BE 198‒277 Caesalpinia sp.
Phellinotus piptadeniae P AU Distinct 3‒6 5‒5.3 ×
3.6‒4.2
BE-E 230‒754 mostly on
Piptadenia spp.
IAC 4365 (paratype) P AU Distinct 3‒5 5.3 × 4.2 BE-E 280‒520 P. gonoachanta
Robledo 1982 P AU Distinct (4‒) 5‒6 5.1 × 4.0 E 250‒500 Piptadenia sp.
URM 80322 AN A Distinct 4‒5 (‒6) 5.1 × 4.1 BE-E 230‒615 Mimosa sp.
URM 80345 P T Distinct (4‒) 5‒6 5.0 × 3.6 E 300‒754 Senegalia sp.
URM 80768 P T Distinct (4‒) 5‒6 5.2 × 3.7 E 268‒504 P. stipulacea
A = applanate, AU = applanate to ungulate, BE = broadly ellipsoid, E = ellipsoid, P = perennial, AN = annual-biannual, U = ungulate, T =
triquetrous. Compilation of variation data about species is presented in bold type.
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 9
FIGURE 2. Phellinotus species. P. neoaridus. (A) holotype on specific host (lenticels of Caesalpinia sp. in detail); (B) applanate to
triquetrous basidioma with a distinct ochre-orange margin (URM80410); (C) ungulate with strongly rimose upper surface and brownish
margin (URM80641) (Bars = 5 cm); (D): isotype context and tubes lacking tissue layer between the tubes, in detail (D*) the black line
near the upper surface (Bars = 1 cm); (E–G) basidiospores, white arrows are indicating the adaxially flattened surface [(E) lactophenol,
(F) KOH+, (G) CB±; Bars = 5 µm]; P. piptadeniae. (H) paratype (IAC4365); (I) context in detail showing the distinct black line in the
paratype; (J) mature triquetrous, developing applanate basidiomata on Piptadenia spp. (URM 80768) (Bars = 5 cm); (L) black line and
tissue layer between the tubes are showing in detail in L and L* (Bars = 1 cm); (M–O) basidiospores, white arrows are indicating the
adaxially flattened surface [(M): lactophenol, (N) KOH+, (O) CB±; Bars = 10 µm]. Photos of basidiomata (A, B, C, D, H, I, J) by Elisandro
Ricardo Drechsler-Santos except for ‘L’ which is by Gerardo Lucio Robledo; photos of spores (E, F, G, M, N, O) by Carlos Salvador-
Montoya.
DRECHSLER-SANTOS ET AL.
10 • Phytotaxa 000 (0) © 2016 Magnolia Press
FIGURE 3. Microscopic elements of Phellinotus species in line drawings. (A, C, E) P. piptadeniae (IAC 4365). (A) hyphal system of
context; (C) skeletal hyphae from trama of tube layer; (E) basidiospores. (B, D, F) P. neoaridus (URM 80362). (B) hyphal system of
context; (D) skeletal hyphae from trama of tube layer; (F) basidiospores. Drawings by Gerardo Lucio Robledo.
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 11
Hymenophore poroid, pores round, some elongated, 3‒6/mm, (90‒)150‒280(‒350) µm wide, reddish yellow to
dark brown; dissepiments entire, (20‒)40‒150(‒300) µm wide. Hyphal system dimitic with skeletal hyphae restricted
to the trama of the tube layer; contextual generative hyphae pale yellow to dark reddish brown, thin to thick-walled,
lumen (3‒)4‒6(‒7) µm wide, regularly simple-septate with ramifications, then becoming sclerified and septa in some
portions of thick-walled hyphae destroyed, sometimes forming secondary septa; trama of tube layer composed of
generative simple-septate and skeletal hyphae; generative hyphae hyaline, pale yellow to fulvous, thin to thick-walled,
(0.5‒)1‒2(‒3) µm wide, forming terminal, dark reddish brown skeletal hyphae, (110‒)232‒533(‒560) × (3‒)4‒6(‒7)
µm, lumen absent to 1–1.5 µm wide, with 1–4 adventitious septa at the narrow to obtuse apex; granular core with few
to many sclereoid hyphae. Basidia not observed. Basidiospores broadly ellipsoid to ellipsoid, 5‒6.5(‒7) × 4‒5(‒5.5)
µm (ave. = 5.8 × 4.7 µm), Q = 1.2‒1.3 (ave.Q = 1.2), adaxially flattened, smooth, thick-walled, yellow, chestnut brown
in KOH, weakly cyanophilous, IKI-.
Habitat and distribution:—Commonly found on living Caesalpinia spp. (Fabaceae) (Drechsler-Santos et al. 2010,
2013, as P. rimosus). Some basidiomata, mostly those that are annual or bi-annual, are found fallen on the ground near
the tree where they presumably developed. Usually, ungulate specimens with many tube layers were found on the tree
trunk as well. This species is widely distributed in the Brazilian semiarid region (Caatinga dry woodlands), from the
states of Alagoas, Bahia, Ceará, Paraíba, Pernambuco, Rio Grande do Norte and Sergipe.
Remarks:—The typical triquetrous to ungulate basidiomata with a black rimose pileus and reddish brown
hymenophore are useful in distinguishing this species. When present, the context of P. neoaridus is weakly zonate
developing one or more dark lines below the rimose upper surface of the pileus and some basidiomata present a very
distinct granular core. Phellinotus piptadeniae is similar, but develops a less rimose pilear surface, a distinct black line
in the context, and smaller basidiospores. Additionally, it grows repeatedly on Piptadenia spp. (Drechsler-Santos et al.
2010, 2013, as Phellinus rimosus).
Material examined:—BRAZIL. Alagoas: São José da Tapera, Fazenda do Sr. Rudá, on living Caesalpinia sp.,
9º32’47” S, 37º33’11” W‒227m alt., 17 June 2008, Drechsler-Santos & al DS017AL (URM 80411, O); ibid, on living
Caesalpinia sp., 9º32’49” S, 37º33’27” W‒224m alt., 17 June 2008, Drechsler-Santos & al. DS009AL (URM 80579);
ibid, on living Caesalpinia sp., 9º32’49” S, 37º33’29” W‒230m alt., 17 June 2008, Drechsler-Santos DS007AL (URM
80410). Ceará: Reriutaba, Serrote de Boqueirão on living Caesalpinia sp., 4º01’13” S, 40º38’43” W‒290m alt., 15
June 2007, Drechsler-Santos DS034CE (URM 80847); ibid, on living Caesalpinia sp., 4º01’15” S, 40º38’41” W‒280m
alt., 15 June 2007, Drechsler-Santos DS033CE (URM 80536). Paraíba: Sumé, Fazenda Almas, on living Caesalpinia
pyramidalis Tul., 7º28’21” S, 36º53’28” W‒686m alt., 10 August 2008, Drechsler-Santos & Rajchenberg DS044PE
(URM 80494). Pernambuco: Cabrobó, Fazenda Mosquito, on living Caesalpinia sp., 08º23’06’’ S, 39º25’30’’ W‒75m
alt., 21 November 2009, Robledo 1974 (CORD); Serra Talhada, Estação Experimental do IPA, on living Caesalpinia
sp. 7º53’49” S, 38º18’12” W‒501m alt., 5 March 2009, Drechsler-Santos & al. DS143PE (URM 80769); ibid, on
living Caesalpinia sp., 7º53’50” S, 38º18’11” W‒560m alt., 5 March 2009, Drechsler-Santos DS151PE (URM 80764);
ibid, on living Caesalpinia sp., 7º53’20” S, 38º18’25” W‒486m alt., 11 March 2008, Drechsler-Santos DS056PE
(URM 80641); ibid, on living Caesalpinia sp., 7º53’47” S, 38º8’12” W‒502m alt., 5 March 2009, Drechsler-Santos
DS140PE (URM 80741); ibid, 7º55’78” S, 38º17’39” W‒504m alt., 12 September 2007, Drechsler-Santos DS22PE
(URM 80299, O). Sergipe: Niterói on living Caesalpinia sp., 9º45’23” S, 37º27’56” W‒57m alt., 16 June 2008,
Drechsler-Santos & al. DS042SE (URM 80422); Poço Redondo, Trilha de Angicos, on living Caesalpinia sp., 9º38’08”
S, 37º40’42” W‒30m alt., 14 June 2008, Drechsler-Santos DS001SE (URM 80419); ibid, on living Caesalpinia sp.,
9º39’08” S, 37º40’42” W‒30m alt., 14 June 2008, Drechsler-Santos & al. DS009SE (URM 80568); Porto da Folha,
on living Caesalpinia sp., 9º54’40” S, 37º16’16” W‒10m alt., 15 June 2008, Drechsler-Santos & al. DS026SE (URM
80577).
Phellinotus piptadeniae (Teixeira) Drechsler-Santos & Robledo comb. nov. (Figs 2H–O, 3B, D, F, Table 2)
≡ Phellinus piptadeniae Teixeira, Bragantia 10: 118, 1950 (IAC, paratype !).
≡ Fomitiporella piptadeniae (Teixeira) Teixeira, Revista Brasileira de Botânica 15(2): 126, 1992.
Mycobank no:—MB805902
Description:—Basidioma annual to bi-annual or perennial, pileate, projecting up to 250 mm, 300 mm wide and 150
mm thick at base, applanate to triquetrous; hard; usually solitary, but also gregarious or many basidiomata widespread
along the substrate; margin applanate, usually distinct from older parts of basidioma, pale brown to brown. Pileus
variable in color, first cream to reddish yellow, becoming olive to black grayish, first azonate, afterwards with sulcate
DRECHSLER-SANTOS ET AL.
12 • Phytotaxa 000 (0) © 2016 Magnolia Press
zones and finally cracked to rimose. Context 20‒60 mm thick, zonate, with a distinct black line separating it in two
parts, pale, fulvous to redish brown, becoming black with KOH, a thin to slightly thick dark brown crust present above
contex. Tubes distinctly stratified, reddish to dark brown, with context layers among the strata of tubes. Hymenophore
poroid, pores round, 3‒6/mm, (100‒)130‒200(‒230) µm wide, reddish yellow to dark brown; dissepiment entire, (40‒
)60‒160(‒200) µm. Hyphal system dimitic with skeletal hyphae restricted to trama of tube layer; contextual generative
hyphae in different stages of development, pale yellow to reddish dark brown, thin to thick-walled, (4‒)5‒8(‒10)
µm wide, always with wide lumen (2‒)2.5‒5(‒8) µm wide, first regularly septate, branched, becoming sclerified
and some portions of thick-walled hyphae sparsely simple-septate; tramal hyphae formed by generative and skeletal
hyphae; generative hyphae simple-septate, hyaline, pale yellow to fulvous, thin to thick-walled in different stages
of development, (1‒)2‒2.5(‒3) µm wide; skeletal hyphae terminal, reddish dark brown, (230‒)280‒470(‒760) µm
long, (3.5‒)5‒8(‒10) µm wide, lumen 1‒2(‒3) µm to solid, some with small aborted ramifications and geniculate
portions at the base or apex, with one to four adventitious septa at the apex, which is narrow to obtuse. Basidia not
found. Basidiospores broadly ellipsoid to ellipsoid, (4.5‒)5‒5.5(‒6) × 3.5‒4.5(‒5) µm (ave.= 5.1 × 3.9 µm), Q =
1.2‒1.4 (ave.Q = 1.3), adaxially flattened, smooth, thick-walled, yellowish, becoming chestnut brown in KOH, weakly
cyanophilous, IKI-.
Habitat and distribution:—Phellinotus piptadeniae is a very common pathogen of living Piptadenia spp.
(Fabaceae), also found on Eugenia L. (1753:470) (Myrtaceae) and on other genera of Fabaceae such as Libidibia
(DC.) Schltdl. (1830: 192), Mimosa L. (1753:516), Pithecellobium Mart. (1837:114), and Senegalia Raf. (1838:119)
(Drechsler-Santos et al. 2010, 2013, Salvador-Montoya et al. 2015, as Phellinus piptadeniae). Most specimens from
Caatinga are annual or bi-annual. Occasionally, basidiomata are large and perennial with many tube layers, such as
the paratype (IAC 4365!) from the SDTFs of Atlantic Forest in Santa Catarina and São Paulo states (Teixeira 1950,
Salvador-Montoya et al. 2015). Additionally, this species is widely distributed in the Brazilian Caatinga dry woodlands
of Alagoas, Bahia, Ceará, Paraíba, Pernambuco, Rio Grande do Norte and Sergipe states and was recently recorded for
lowland SDTF of northwestern Peru (Salvador-Montoya et al. 2015).
Remarks:—This species is morphologically variable, but the slightly rimose pileus, fulvous to reddish brown
context and a longitudinal dark line in the middle of the basidioma context are diagnostic (Drechsler-Santos et al.
2010, Salvador-Montoya et al. 2015). Phellinotus piptadeniae presents slightly smaller basidiospores (ave.= 5.1 ×
3.9 µm versus 5.8 × 4.7 µm) than those of P. neoaridus. Additionally, it is repeatedly found on living Piptadenia spp.
(Drechsler-Santos et al. 2010), not yet recorded on Caesalpinia spp.
Material examined:—BRAZIL. Alagoas: São José da Tapera, Fazenda do Sr. Rudá, on living Mimosa sp., 9º32’45”
S, 37º33’12” W‒231m alt., 17 June 2008, Drechsler-Santos DS16AL (URM 80414). Bahia: Senhor do Bonfim, Serra
do Santana, Fazenda Passaginha, Vasconcellos-Neto J.R.T. 195 (HUEFS 133884, O). Ceará: Reriutaba, Boqueirão, on
living Piptadenia stipulacea (Benth.) Ducke (1930:126), 4º01’30” S, 40º38’21” W‒197m alt., 14 June 2007, Drechsler-
Santos DS23CE (URM 80466, O). Paraíba: Sumé, Fazenda Almas, on living Mimosa ophthalmocentra Mart. ex Benth.
(1875:415), 7º28’21” S, 36º53’28” W‒686m alt., 11 August 2008, Drechsler-Santos & Rajchenberg DS52PB (URM
80498). Pernambuco: Cabrobó, Fazenda Mosquito, on living Mimosa sp., 21 September 2009, Robledo 1982 (CORD);
Caruaru, Estação Experimental do IPA, on living Mimosa sp., 8º13’50” S, 35º55’14” W‒569m alt., 6 August 2006,
Drechsler-Santos DS163PE (URM 80766, O); ibid, on living Senegalia sp., 8º13’51” S, 35º55’13” W‒558m alt., 10
December 2008, Drechsler-Santos & al. DS128PE (URM 80361); ibid, on living Mimosa sp., 8º13’53” S, 35º55’12”
W‒562m alt., 10 December 2008, Drechsler-Santos & al. DS109PE (URM 80322); ibid, on living Senegalia sp.
(CB01), 10 December 2008, Drechsler-Santos & al. DS110PE (URM 80345); Parque Nacional do Catimbau, Barro
Branco, on living Piptadenia moniliformis Benth. (1841:339‒340), 8º30’0” S, 37º18’57” W‒818m alt., 9 February
2007, Drechsler-Santos & al. DS158 (URM 80884); ibid, Breu, on living P. stipulacea, 8º30’29” S, 37º16’58” W‒
964m alt., 22 June 2007, Drechsler-Santos DS240 (URM 80854, O); ibid, Casa da Farinha, on living P. stipulacea,
8º33’38” S, 37º13’89” W‒839m alt., 31 October 2007, Drechsler-Santos & al. DS280 (URM 80602); ibid, Cerca
de Pedra, on living P. moniliformis, 8º29’04” S, 37º19’24” W‒723m alt., 9 February 2007, Drechsler-Santos & al.
DS152 (URM 80694); ibid, Dor de Dente, on living P. stipulacea, 8º30’33” S, 37º16’59” W‒958m alt., 23 July 2007,
Drechsler-Santos & al. DS253 (URM 80737); ibid, Morro do Cachorro, on living Pithecollobium polycephallum
Benth. (1844:219), 8º34’28” S, 37º14’47” W‒751m alt., 5 August 2006, Drechsler-Santos DS64 (URM 80667, O);
ibid, Muquém, on living P. moniliformis, 8º29’42” S, 37º19’03” W‒795m alt., 29 September 2006, Drechsler-Santos
DS85 (URM 80713); ibid, on living P. stipulacea, 8º30’10” S, 37º17’76” W‒921m alt., 25 January 2008, Drechsler-
Santos & al. DS300 (URM 80595); ibid, Pedra Solteira, on living P. moniliformis, 8º29’42” S, 37º19’03” W‒795m alt.,
27 May 2006, Drechsler-Santos DS003 (URM 80825); ibid, on living P. stipulacea, 8º32’52” S, 37º14’38” W‒922m
alt., 15 April 2007, Drechsler-Santos & al. DS207 (URM 80697); ibid, Trilha da Batinga, on living P. stipulacea,
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 13
8º30’47” S, 37º16’57” W‒957m alt., 28 September 2006, Drechsler-Santos & al. DS76 (URM 80730); ibid, Trilha
das Torres, on living P. stipulacea, 8º34’20” S, 37º14’44” W‒764m alt., 8 July 2006, Drechsler-Santos DS101 (URM
80662, O); ibid, Trilha do Pereira, on dead Eugenia rostrifolia D. Legrand (1957:2), 8º30’37” S, 37º14’39” W‒943m
alt., 10 July 2006, Drechsler-Santos & al. DS133 (URM 80890); Petrolina, Embrapa Semi-árido, on living Mimosa
arenosa (Willd.) Poir. (1810:66), 9º04’22” S, 40º01’91” W‒386m alt., 17 April 2008, Drechsler-Santos DS6PETRO
(URM 80436, HTSA, O); Serra Talhada, Estação Experimental do IPA, on living P. stipulacea, 7º53’27” S, 38º18’22”
W‒503m alt., 11 March 2008, Drechsler-Santos DS72PE (URM 80631); ibid, on living Mimosa sp., 11 March 2008,
Drechsler-Santos & al. DS74PE (URM 80317); ibid, on living Piptadenia sp., 7º54’01” S, 38º18’05” W‒513m alt.,
5 March 2009, Drechsler-Santos DS139PE (URM 80768, O); ibid, on living Mimosa sp., 7º53’29” S, 38º18’17” W‒
490m alt., 9 July 2008, Drechsler-Santos & al. DS97PE (URM 80360, O). Rio Grande do Norte: Estação Ecológica do
Seridó-ESEC, 19 February 2007, Silva & al. Sn (URM 80883). Sergipe: Niterói, 9º45’20” S, 37º27’55” W‒65m alt.,
16 June 2008, Drechsler-Santos & al. DS40SE (URM 80567).
Discussion
Phellinotus and its phylogenetic affinities
Phellinotus is considered a pathogenic polypore, common on living members of the family Fabaceae, and so far
endemic to the Neotropics. Within the ‘phellinotus clade’, Phellinotus is closely related, both phylogenetically and
morphologically, to Arambarria, Inocutis, Fomitiporella and some other taxonomically unresolved lineages (Fig. 1,
Table 1). Phellinotus develops skeletal hyphae in the trama of the tube layer whereas Arambarria and Inocutis lack
skeletal hyphae entirely. Fomitiporella differs from Phellinotus by its saprotrophic habit, resupinate basidiomata and
skeletal hyphae that occur throughout the basidioma (Murrill 1907, Teixeira 1994, Zhou 2014b). Inocutis dryophila,
the closest lineage in our phylogeny, is distributed in the Holartic region, has ellipsoid to ovoid, thick-walled and
brownish basidiospores, as well as a monomitic hyphal system throughout the basidioma; it also grows on living oaks
(Ryvarden 2005). Fulvifomes chinensis and F. inermis also present broadly ellipsoid, fairly thick-walled, yellowish
and cyanophilous basidiospores (Dai 2010); on the other hand, they have effused-reflexed and resupinate basidiomata,
respectively, besides a dimitic hyphal system both in the context and trama of the tubes. Their phylogenetic position is
still unresolved (Zhou 2014a). Inonotus tenuissimus was described from China as having ellipsoid, thick-walled and
yellowish basidiospores and a monomitic to dimitic hyphal system with “skeletal-like” hyphae in the trama of tube
layer (Yu et al. 2013), which could be interpreted as a similar hyphal system to Phellinotus. However, the resupinate
and adnate annual basidiomata and absence of cyanophilous reaction in the basidiospores of I. tenuissimus are quite
different from the new genus. Isolates CIEFAPcc107 and cc88 from Patagonia (named as Hymenochaetaceae sp.)
are known only from mycelium isolates of wood and their basidiomata have not been recorded; they form a distinct
phylogenetic species (Rajchenberg et al. 2015).
Phellinotus is also phylogenetically close to Fulvifomes s.s. and Phylloporia. This group of related genera,
previously noted by Wagner & Fischer (2002), Larsson et al. (2006), Zhou (2014a), Rajchenberg et al. (2015) and Zhou
et al. (2015), named here as the ‘phellinotus clade’, deserves special taxonomic recognition. This is a group of poroid
Hymenochaetaceae taxa with thick-walled and colored basidiospores and without setae or setal hyphae. Aditionally, the
adaxially flattened basidiospores seems to be an important taxonomic feature and probably phylogenetically significant
for the ‘phellinotus clade’. Actually, despite Phellinotus, in our morphological analysis of types of Arrambaria (A.
destruens), Fomitiporella (F. umbrinella), Fulvifomes (Pyropolyporus robiniae), and Phylloporia (P. parasitica)
we were able to observe adaxially flattened basidiospores. Type specimens of Fulvifomes chinensis, F. inermis and
Inonotus tenuissimus should be revised to confirm this character. This is a feature usually not checked and not found
in the literature, except by Kotlaba & Pouzar (1978, 1979), who mentioned it in the description of Phellinus robiniae
(Murrill) A. Ames (1913:246), P. rimsosus s.s., P. coffeatoporus Kotl. & Pouzar (1979: 259), and P. resinaceus Kotl.
& Pouzar (1979: 261).
Fulvifomes s.s. (F. rimosus group excluding F. chinensis and F. inermis) is macro- and micro- morphologically
similar to Phellinotus, but distinguishable mainly by the shape of its basidiospores and hyphal system (Table 1). The
shape of the basidiospore is an important feature, as recognized by Murrill (1914), who segregated Fulvifomes from
Pyropolyporus Murrill (1903:109) based on Pyropolyporus robiniae [≡ Fulvifomes robiniae (Murrill) Murrill (1914:49)]
as genus type. The lectotype of Phellinus robiniae presents subglobose, 5.5‒6 × 4.5‒5(‒5.5) µm basidiospores.
Aurificaria is characterized by basidiospores that turn dark olivaceous green in KOH and by a monomitic hyphal
DRECHSLER-SANTOS ET AL.
14 • Phytotaxa 000 (0) © 2016 Magnolia Press
system (Ryvarden 2004). Phylogenetic studies by Larsson et al. (2006) and Zhou (2014c), based on ITS and nLSU,
show that Aurificaria is a basal sister clade of Fulvifomes s.s. Zhou (2014c) proposed its synonymy with Fulvifomes,
which would expand the concept of Fulvifomes by including monomitic and dimitic species with annual or perennial
basidiomata and basidiospores that can turn brown or green in KOH.
Phylloporia species are mostly parasitic on lianas and roots of shrubs and trees, which are difficult to identify
in the Neotropics (Ryvarden 2004). As a result, the extent of host specialization of Phylloporia species is unknown.
Phylloporia and Phellinotus share the black line in the context; however, the former is distinguished by its smaller,
yellow basidiospores that remain slightly yellowish in KOH (Ryvarden 2004).
The occurrence of different types of hyphal systems in the same basidioma is well documented and has been
used to distinguish taxa (Niemelä et al. 2001, Tian et al. 2013). The Inonotus linteus (Berk. & M.A. Curtis) Teixeira
(1992:126) species-complex, like Phellinotus species, develops perennial basidiomata with skeletal hyphae restricted
to the trama of the tube layer (Tian et al. 2013, Vlasák et al. 2013). However, it differs from Phellinotus by the presence
of hymenial setae and is not phylogenetically related. Phylogenetic studies confirmed its relationship with Inonotus
(Wagner & Fischer 2002), and the closely related recently described genera Sanghuangporus Sheng H. Wu, L.W.
Zhou (2015:6) & Y.C. Dai and Tropicoporus L.W. Zhou, Y.C. Dai & Sheng H. Wu (2015:7)(Zhou et al. 2015), are also
positioned outside of the ‘phellinotus clade’. Indeed, the sequences available in GenBank, identified as Inonotus linteus
(Tropicoporus) should be taxonomically re-evaluated, as some vouchers are actually placed in the Sanghuangporus
clade (e.g., vouchers F915611 and MUCL74139) (Fig. 1).
The current phylogenetic scenario shows that the diversity of the group is higher than expected and the incorporation
of specimens and taxa from other unexplored regions of the world into phylogenetic studies is necessary to attain a
better understanding of the diversity and natural classification of Hymenochaetaceae.
Phellinotus and its species
Phellinus piptadeniae was proposed by Teixeira (1950) and described as host specific on living Piptadenia communis
Benth. [=Piptadenia gonoacantha (Mart.) J.F. Macbr. (1919:17)]. He described it as having a large basidioma with
a smooth pilear surface that becomes rimose at maturity, a golden brown and lustrous context with a distinct black
line, and stratified tubes forming 4‒5 pores/mm on the hymenophore. Regarding particular microscopic characters,
the hyphal construction of the basidioma deserves special attention. Teixeira (1950) described two types of hyphae
in the context, branched or not, with walls not much thickened and simple-septa present in the unbranched hyphae.
He did not mention the specific type of hyphal system present. Later, he transferred this species to Fomitiporella as
F. piptadeniae (Teixeira) Teixeira (1992:126) (Teixeira 1992), again without describing the hyphal system. Later, he
described Fomitiporella as dimitic and closely related to Phellinus and Phylloporia (Teixeira 1994).
According to Ryvarden (2004), Phellinus piptadeniae is closely related to Phellinus rimosus; moreover, he
suggested that molecular work would be necessary to determine if P. piptadeniae was an immature stage of P. rimosus
or not. Phellinus piptadeniae is morphologically and ecologically a well-delimited species (Teixeira 1950, Drechsler-
Santos et al. 2010), whereas Phellinus rimosus is considered to be a taxonomic complex of three or more cryptic
species, namely, P. badius (Berk. ex Cooke) G. Cunn. (1965:273), P. rimosus s.s. and P. robiniae (Kotlaba & Pouzar
1978). Interestingly, most of these species seem to be host-specific and geographically restricted (Kotlaba & Pouzar
1978, 1979). Species in the P. rimosus complex are characterized by having woody, pileate basidiomata with a cracked
to strongly rimose pileus, a lack of setae, and ellipsoid, thick-walled, chestnut to rusty brown basidiospores (Fiasson
& Niemelä 1984). Fulvifomes has a dimitic hyphal system and globose to subglobose basidiospores (Table 1).
Key to Phellinus s.l. and Inonotus s.l. and related poroid/hydnoid genera of Hymenochaetaceae
1. Basidiospores hyaline and slightly to strongly dextrinoid .................................................................................................................2
1. Basidiospores hyaline to chestnut brown and not dextrinoid .............................................................................................................4
2. Basidiospores ellipsoid, hyphal system monomitic, setae (hymenial or tramal) present ..................................................Mensularia
2. Basidiospores globose, hyphal system monomitic or dimitic, if present, setae only hymenial .........................................................3
3. Basidiomata perennial, resupinate to pileate, setae absent, occasionally present ............................................................ Fomitiporia
3. Basidiomata annual, pileate, hymenial setae present ................................................................................................ Pseudoinonotus
4. Setae (any type) present ......................................................................................................................................................................5
4. Setae (any type) absent .....................................................................................................................................................................17
5. Asterosetae present .......................................................................................................................................................Asterodon Pat.
5. Asterosetae absent, hymenial setae and/or setal hyphae present ........................................................................................................6
6. Hyphal system monomitic throughout ...............................................................................................................................................7
6. Hyphal system dimitic, at least in the trama of tube layer, or trimitic .............................................................................................11
PHELLINOTUS, A NEW NEOTROPICAL GENUS Phytotaxa 000 (0) © 2016 Magnolia Press • 15
7. Setal hyphae present ...........................................................................................................................................................................8
7. Setal hyphae absent ............................................................................................................................................................................9
8. Basidiospores hyaline, thin-walled, hymenial setae absent ........................................................................................... Phellinidium
8. Basidiospores yellow to brown, thick-walled, hymenial setae absent or present ...........................................................Inonotus s.s.
9. Basidiomata sessile pileate to substipitate, basidiospores thin-walled ............................................................................ Inonotus s.l.
9. Basidiomata sessile resupinate/pileate, basidiospores thick-walled .................................................................................................10
10. Basidiomata stipitate, sub-stipitate or centrally sessile, basidiospores ellipsoid to globose, tramal to hymenial setae present ..........
......................................................................................................................................................................................Onnia P. Karst
10. Basidiomata sessile, basidiospores cylindrical, hymenial setae present ...............................Cylindrosporus L.W. Zhou & Y.C. Dai
11. Basidiospores obclavate to cylindric, hyphal system dimitic-trimitic ............................................................................... Phellopilus
11. Basidiopores ellipsoid, broadly ellipsoid to globose-subglobose, hyphal system dimitic ...............................................................12
12. Basidiospores thin-walled, basidiospores ellipsoid/allantoid/cylindrical, generative hyphae with crystals .....................Fuscoporia
12. Basidiospores slightly thick- to thick- walled, basidiospores ellipsoid/to subglobose-globose, generative hyphae without crystals .
..........................................................................................................................................................................................................13
13. Hyphal system mono-dimitic, contex monomitic, trama of tube layer dimitic ................................................................................14
13. Hyphal system dimitic throughout ...................................................................................................................................................15
14. Basidiomata annual to perennial, resupinate to pileate, basidiospores when ellipsoid mostly < 3.5 μm wide, exclusively in tropical
zone .................................................................................................................................................................................Tropicoporus
14. Basidiomota perennial, resupinate, effused-reflexed to pileate, basidiospores when ellipsoid mostly > 3.5 μm wide, in boreal,
temperate, subtropical to tropical zones .................................................................................................................. Sanghuangporus
15. Tramal setae penetrating into hymenium ........................................................................................................................ Phellinopsis
15. Tramal setae absent ..........................................................................................................................................................................16
16. On coniferous wood ......................................................................................................................................................Porodaedalea
16. On hardwood ..................................................................................................................................................................Phellinus s.s.
17. Basidiospores thin-walled ...............................................................................................................................................................18
17. Basidiospores thick-walled ..............................................................................................................................................................20
18. Basidiospores cylindrical to obclavate ...................................................................................................................... Nothophellinus
18. Basidiospores subglobose to ellipsoid .............................................................................................................................................19
19. Basidiomata resupinate, basidiospores ellipsoid, on coniferous wood ........................................................................... Inonotopsis
19. Basidiomata pileate/substipitate, basidiospores subglobose, on deciduous wood .........................................................Pyrrhoderma
20. Hyphal system monomitic ................................................................................................................................................................21
20. Hyphal system dimitic, at least in the trama of tube layer ...............................................................................................................25
21. Basidiospores ellipsoid smaller than 5 µm, hyaline in KOH ...........................................................................................Phylloporia
21. Basidiospores usually larger than 5 µm ...........................................................................................................................................22
22. Basidiospores yellowish, becoming dark olivaceous green in KOH ................................................................................ Aurificaria
22. Basidiospores do not become dark olivaceous green in KOH .........................................................................................................23
23. Basidiospores yellowish to yellowish brown, becoming chestnut brown in KOH, thick-walled and adaxially flattened ............. 24
23. Basidiospores without this combination of characters .................................................................................................. Inonotus s.l.
24. Granular core present at the base of the context ................................................................................................................... Inocutis
24. Granular core absent at the base of the context .............................................................................................................. Arambarria
25. Basidiospores usually shorter than 5 µm, hyaline in KOH ............................................................................................. Phylloporia
25. Basidiospores usually larger than 5 µm, becoming chestnut brown in KOH ................................................................................. 26
26. Basidiospores globose to subglobose ...............................................................................................................................Fulvifomes
26. Basidiospores ellipsoid to broadly ellipsoid ....................................................................................................................................27
27. Skeletal hyphae restricted to the trama of tube layer ........................................................................................................Phellinotus
27. Skeletal hyphae found throughout the basidiomata .........................................................................................................................28
28. Basidiomata resupinate ..................................................................................................................................................Fomitiporella
28. Basidiomata pileate ........................................................................................................................................................ Phellinus s.l.
Acknowledgements
The authors thank Aristóteles Góes-Neto, Clarice Loguercio-Leite, Carlos Urcelay, Cony Decock, Domingos Cardoso,
Gladstone Silva, Leonor Costa Maia, and Carlos Alberto Salvador Montoya for their valuable contributions to
this work. Curators of herbaria mentioned here are thanked for the loan of exsiccates. The managers of the study
areas are thanked for permission to sample collections. This research was supported by the Conselho Nacional de
Desenvolvimento Científico e Tecnológico – CNPq-Brazil (Universal 478973/2006-3 and 479961/2007-7, Pesquisa
em Biodiversidade do Semi-árido 010105.00/2004/PPBio) and the Coordenação de Aperfeiçoamento Pessoal de Nível
Superior–CAPES-Brazil and Ministerio de Ciencia, Tecnología e Innovación Productiva–MINCYT-Argentina (Project
161/09 - BR/08/13). CNPq, MINCYT and CAPES supported the first author’s PhD (modalities: GD 141072/2006-7
and SWE 201847/2008-6) and Post-doctoral scholarships (CAPES-MINCYT and CAPES/REUNI). GLR and MR
are members of the Research Career of CONICET (Argentina). MAR thanks CAPES (PNPD Institucional 2011 -
23038.007790/2011-93) for Postdoctoral scholarship.
DRECHSLER-SANTOS ET AL.
16 • Phytotaxa 000 (0) © 2016 Magnolia Press
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APPENDIX 1. List of species, sources, localities, and GenBank accession numbers of taxa used in this study.
Taxon; Geographic origin; Collection reference; Substrate;
GenBank accession.
nLSU sequences
Arambarria destruens Rajchenb. & Pildain; Argentina/Chubut;
CIEFAPcc 347; Diostea juncea; KP347523.
Cylindrosporus flavidus L.W. Zhou; China, Da 13213; on
fallen angiosperm wood; KP875561.
Fomitiporella sp.; Ethiopia; Oe5; JQ910908. Fomitiporella
sp.; Ethiopia; Oe6; JQ910909.
Fomitiporia hartigii (Allesch. & Schnabl) Fiasson &
Niemel; Europe/Estonia; MUCL 53551; JX093833.
F. mediterránea M. Fischer; Europe/Italy; MUCL
38514; living branch; AY618201. F. punctata (Fr.)
Murrill; Europe/Estonia; MUCL 53548; JX093834. F.
robusta (P. Karsten) Fiasson & Niemel; Europe/Czech
Republic; MUCL 51327; GU461993.
Fulvifomes chinensis (Pilát) Y.C. Dai; China/ Zhashui; LWZ
20130916-3; on dead standing angiosperm; KJ787809.
F. chinensis; China/ Weishan; LWZ 20130713-7; on
fallen angiosperm trunk; KJ787808. F. fastuosus (Lév.)
Bondartseva & S. Herrera; Philippines; CBS 213.36;
Gliricidia sepium; AY059057. F. indicus (Massee)
L.W. Zhou; Zimbabwe/Midlands; O 25034; KC879259.
F. indicus (Massee) L.W. Zhou; China/Ningming;
Yuan 5932; on living tree of Bombax ceiba; JX866777.
F. inermis (Ellis & Everh.) Y.C. Dai; China/Qimen;
LWZ 20130810-3; on fallen angiosperm branch;
KJ787812. F. inermis; China/Qimen; LWZ 20130809-
8; on living angiosperm tree; KJ787811. F. merrillii
(Murrill) Baltazar & Gibertoni; Taiwan; JX484002. F.
nilgheriensis (Mont.) Bondartseva & S. Herrera; USA;
CBS 209.36; on dead deciduous wood; AY059023. F.
rimosus (Berk.) Fiasson & Niemel; Taiwan; JX484003.
F. robiniae (Murrill) Murrill; USA; CBS 211.36;
Robinia pseudo-acacia; AY059038.
Fuscoporia atlantica Motato-Vásquez, Pires & Gugliotta;
Brazil; MV230; KP058517. F. gilva (Schwein.) T.
Wagner & M. Fisch.; Brazil; RP 17; KP859305. F.
senex (Nees & Mont.) Ghob.-Nejh.; KUC20110922-13;
JX463652. F. torulosa (Pers.) T. Wagner & M. Fisch.;
Csezh Republic; 759; AM269865.
Hymenochaetaceae sp.; Argentina/Chubut; CIEFAPcc88;
Austrocedrus chilensis; KP347524. Hymenochaetaceae
ITS sequences
Arambarria destruens Rajchenb. & Pildain; Argentina/Chubut;
CIEFAPcc 347; Diostea juncea; KP347538.
Cylindrosporus flavidus L.W. Zhou; China, Da 13213; on fallen
angiosperm wood; KP875564.
Fomitiporella sp.; Ethiopia; Oe5; JF895466. Fomitiporella sp.;
Ethiopia; Oe6; JF895467.
Fomitiporia hartigii (Allesch. & Schnabl) Fiasson & Niemel;
Europe/Estonia; MUCL 53551; JX093789. F.
mediterránea M. Fischer; Europe/Italy; MUCL 38514;
living branch; GU461953. F. punctata (Fr.) Murrill;
Europe/Estonia; MUCL 53548; JX093790. F. robusta
(P. Karsten) Fiasson & Niemel; Europe/Czech Republic;
MUCL 51327; GU461949.
Fulvifomes chinensis (Pilát) Y.C. Dai; China/ Zhashui; LWZ
20130916-3; on dead standing angiosperm; KJ787818.
F. chinensis; China/ Weishan; LWZ 20130713-7; on
fallen angiosperm trunk; KJ787817. F. fastuosus (Lév.)
Bondartseva & S. Herrera; Philippines; CBS 213.36;
Gliricidia sepium; AY558615. F. inermis (Ellis &
Everh.) Y.C. Dai; China/Qimen; LWZ 20130810-3; on
fallen angiosperm branch; KJ787821. F. inermis; China/
Qimen; LWZ 20130810-8; on living angiosperm tree;
KJ787820. F. indicus (Massee) L.W. Zhou; Zimbabwe/
Midlands; O 25034; KC879262. F. indicus (Massee)
L.W. Zhou; China/Ningming; Yuan 5932; on living
tree of Bombax ceiba; KC879261. F. merrillii
(Murrill) Baltazar & Gibertoni; Taiwan; JX484013. F.
nilgheriensis (Mont.) Bondartseva & S. Herrera; USA;
CBS 209.36; on dead deciduous wood; AY558633. F.
rimosus (Berk.) Fiasson & Niemel; Taiwan; JX484016.
F. robiniae (Murrill) Murrill; USA; CBS 211.36;
Robinia pseudo-acacia; AY558646.
Fuscoporia atlantica Motato-Vásquez, Pires & Gugliotta;
Brazil; MV230; KP058515. F. gilva (Schwein.) T.
Wagner & M. Fisch.; Brazil; RP 17; KP859295. F.
senex (Nees & Mont.) Ghob.-Nejh.; KUC20110922-13;
JX463658. F. torulosa (Pers.) T. Wagner & M. Fisch.;
Csezh Republic; 759; AM269803.
Hymenochaetaceae sp.; Argentina/Chubut; CIEFAPcc 88;
Austrocedrus chilensis; KP347536. Hymenochaetaceae
