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Morphological and Phylogenetic Reevaluation of the
Genera Mycobonia and Pseudofavolus
(Polyporaceae)
Melissa Palacio ( melissapalacio@gmail.com )
Universidade Federal do Rio Grande do Sul https://orcid.org/0000-0003-2890-5189
Mauro Westphalen
Universidade Federal do Rio Grande do Sul
Yuan Yuan
Beijing Forestry University
Yingda Wu
China Fire and Rescue Institute
Rosa Mara Borges Da Silveira
Universidade Federal do Rio Grande do Sul
Research Article
Keywords: polypores, core polyporoid clade, phylogeny, hymenophore conguration
Posted Date: September 30th, 2021
DOI: https://doi.org/10.21203/rs.3.rs-925842/v1
License: This work is licensed under a Creative Commons Attribution 4.0 International License.
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Abstract
Mycobonia
and
Pseudofavolus
(Polyporales, Basidiomycota) are polyporoid genera with tropical and
subtropical distribution. Both genera are morphologically similar in presenting abelliform to conchate
subsessile basidiomata, with a dimitic hyphal system, consisting of clamped generative hyphae and
skeleto-binding hyphae that produce large basidiospores with smooth, thin walls. However, while
Pseudofavolus
species present a poroid hymenophore, in
Mycobonia
it is stereoid with hyphal pegs that
resemble thin teeth.
Mycobonia
and
Pseudofavolus
have a controversial taxonomy, and the phylogenetic
relationships between their species have yet to be assessed. For this reason, we performed molecular
phylogenetic analyses on specimens of
Mycobonia
and
Pseudofavolus
from both the Neotropics and
Asia, using internal transcribed spacers (ITS), the large subunit of nuclear rDNA (nc LSU rDNA), and also
the genes encoding the second largest subunit of RNA polymerase II (
RPB2
). Furthermore, in order to
develop an evolutionary analysis of the hymenophore conguration, we performed stochastic character
mapping of ancestral states for the hymenophore type presented in
Polyporus
s.l.
Our study revealed that
Pseudofavolus
is an articial group and its species actually nest in a clade within
Mycobonia
. Therefore,
in order to establish a monophyletic group, based upon priority of publication, we re-circunscribed
Mycobonia
to encompass both stereoid and poroid hymenophore species. Two new combinations are
presented from the Neotropics:
Mycobonia cucullata
and
M. miquelii
. A new species from tropical Asia,
M. yuchengii
, is also described. We presente a summary of stochastic mapping of ancestral states
estimates of hymenophore type in
Polyporus
s.l
. The ancestral state for
Mycobonia
clade is estimated to
have angular pores.
Introduction
Mycobonia
Pat. and
Pseudofavolus
Pat. are genera of wood-decaying Basidiomycota belonging to the
family Polyporaceae with tropical to subtropical distribution. Both genera are morphologically similar in
presenting abelliform to conchate subsessile basidiomata, a dimitic hyphal system with clamped
generative hyphae, skeleto-binding hyphae and large, smooth basidiospores with thin and smooth walls
(Corner 1984).
Macroscopically, the shape, color and consistency of the basidiomata are very similar in both genera;
however, while
Pseudofavolus
includes species with poroid hymenophore,
Mycobonia
presents stereoid
hymenophore with hyphal pegs (Corner 1984; Julich 1976).
Currently, two species are recognized in
Mycobonia
,
M. brunneoleuca
and
M. ava
(type species), and are
distributed throughout the Americas in tropical and subtropical areas, though they differ in their
basidiospores differ in shape, size and altitudinal occurrence (Julich 1976; Gerlach and Loguercio-Leite
2011). Currently nine species are accepted in the
Pseudofavolus
genus, with tropical and subtropical
worldwide distribution (Ryvarden and Johansen 1981; Ryvarden 2016). Specically, four of them were
originally described from the Neotropics:
P. cucullatus
(Mont.) Pat.,
P. miquelii
(Mont.) Pat. (type species),
P. nigrus
Ryvarden, and
P. orinocensis
(Pat. and Gaillard) Ryvarden. Additionally,
P. cucullatus
, which was
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originally described from Cuba, has subsequently been recorded from several areas of subtropical and
tropical Asia (Dai 2007, 2012; Dai et al. 2011; Cui et al. 2019; Wu et al. 2020).
The taxonomy of
Mycobonia
and
Pseudofavolus
is controversial. Due to the stereoid hymenophore,
Mycobonia
has been included in several families, such as Corticiaceae, Mycoboniaceae, Stereaceae and
Thelephoraceae (Corner 1984; Krueger 2002). However, in several studies, the anities of
Mycobonia
and
Pseudofavolus
with
Polyporus s.l.
were described and discussed based on morphological characteristics
(Singer 1951, 1986; Corner 1984). These similarities were subsequently corroborated by phylogenetic
studies (Kruguer 2002; Krueger and Gargas 2004, Motato-Vásquez et al. 2018).
Mycobonia
and
Pseudofavolus
have also been treated as synonyms of
Polyporus
(Silveira and Wright 2005; Nakasone
2015). Nevertheless, phylogenetic studies have placed
Mycobonia
next to
Pseudofavolus
, and in distant
relation to
Polyporus s.s
. (Krüger and Gargas 2004; Motato-Vasquez et al. 2018), thus representing
different genera in the core polyporoid clade. In the present study, we aim to investigate the relationships
between
Mycobonia
and
Pseudofavlus
based on morphological and molecular evidence. In addition, we
compare the Asian specimens of
P. cucullatus
to neotropical samples in order to determine if they are in
fact conspecic. Finally, considering the traditional taxonomic value of hymenophore differentiation, we
reconstruct the ancestral state for the hymenophore conguration of
Polyporus s.l
.
Material And Methods
Specimens and morphological studies
The studied specimens are deposited in ICN, BJFC, and SP herbaria. Herbarium acronyms follow Thiers
(continuously updated, http://sweetgum.nybg.org/science/ih/). Freehand cross sections of dried
materials were mounted on slides and observed under the microscope in Melzer’s reagent, 5% KOH
and/or 1% phloxine, cresyl blue and/or cotton blue (CB). To observe hyphal systems, small pieces of
basidiomata were kept in 3% NaOH solution at 45 ºC for about 2 h; then, pieces were dissected under a
stereomicroscope (Decock et al. 2013). Basidiospores were measured (n = 40) in Melzer’s reagent. The
following abbreviations were also used: IKI– = inamyloid and non-dextrinoid, CB+/– =
cyanophilous/acyanophilous, m = arithmetic mean and Q = the ratio of length/width of basidiospores. We
followed Stalpers (1996) and the Stalpers database (http://www.cbs.knaw. nl/russulales/) for
basidiospore shape terminology.
DNA extraction and sequencing
Following the manufacturer's protocols, DNA was extracted from dried specimens using CTAB rapid plant
genome extraction kit-DN14 (Aidlab Biotechnologies Co., Ltd., Beijing, China) and FH plant DNA kit II
(Demeter Biotech Co., Ltd., Beijing, China). Primer pairs ITS4/ITS5 (White et al. 1990), LR0R/LR7 (Vilgalys
and Hester 1990) and fRPB2-5F/bRPB2-7.1R (Matheny 2005; Frøslev et al. 2005) were used to amplify
the ITS, nc LSU rDNA and
RPB2
regions, respectively, by a qualitative simplex Polymerase Chain Reaction
(PCR). The parameters of the PCR for each region were followed as described by Dentinger et al. (2010),
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Vilgalys and Hester (1990), Frøslev et al. (2005) and Matheny (2005), respectively. The PCR products
were puried and sequenced with the same primers at Beijing Genomics Institute in China.
Phylogenetic analyses
Chromatograms were assembled and manually edited using Geneious v. 7.1.9
(http://www.geneious.com). The newly generated ITS, nc LSU rDNA and
RPB2
sequences were deposited
in GenBank and were later combined with additional sequences retrieved from GenBank to compose the
entire dataset (Table 1). The ITS, nc LSU rDNA and
RPB2
matrices were individually aligned using MAFFT
v.7 (Katoh and Standley 2013) under the Auto strategy, then inspected and edited using Aliview (Larsson
2014). We used PartitionFinder v.2 (Lanfear et al. 2017) to estimate the best-t partitioning strategy and
the best-t model of nucleotide evolution with the following settings: branch lengths = linked, models = mr
bayes, model selection = AICc and search = greedy. Two distinct analyses were performed: Bayesian
Inference (BI) and Maximum Likelihood (ML). Bayesian Inference analysis was conducted using MrBayes
3.2 (Ronquist et al., 2012) with two independent runs, each one with four chains and starting from
random trees. The runs performed 20 million generations, and trees were sampled every 1,000th
generation. Of the sampled trees, 25% were discarded as burn-in, while the remainder were used for
calculating a consensus tree and Bayesian Posterior Probabilities (BPP).
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Table 1
List of species, collections, geographic origin and GenBank accession numbers for the ITS, nc LSU rDNA,
mtSSU and RPB2 sequences used in the phylogenetic analyses in this study. New sequences generated
from this study are marked in bold.
Taxon Voucher Country GenBank accession number
ITS nc LSU
rDNA
RPB2
Atroporus diabolicus
DS1266 Brazil KY631757 KY631768 —
Atroporus rufoatratus
DS816 Brazil KY631759 KY631770 KY744947
Bresadolia craterella
TENN59383 Ecuador AJ487944 —
Bresadolia paradoxa
MV23 Brazil KY777230 KY777235 —
Bresadolia paradoxa
Robledo1958 Argentina KY777233 KY777237 —
Cerioporus
squamosus
AFTOLID704 DQ267123 AY629320 DQ408120
Datronia mollis
RLG6304sp USA JN165002 JN164791 JN164872
Datronia stereoides
Holonen Finland KC415179 KC415196 KC415202
Datroniella
mellanocarpa
Cui10646 China KC415186 KC415194 KC415201
Datroniella scutellata
RLG9584T USA JN165004 JN164792 JN164873
Dichomitus
campestris
O103769 Norway — AJ487512 —
Dichomitus
sp. IFP14643 China KX832053 KX832062 —
Echinochaete
brachypora
TFMF24996 Japan AB462321 AB462309 —
Echinochaete
russiceps
TFMF15716 Japan AB462310 AB368065 AB368123
Echinochaete
sp. MN272 Ecuador AF518754
Favolus brasiliensis
Kellermann
s.n. Brazil MN648682 MN648708 —
Favolus emerici
WD2379 Japan AB587628 AB587619 AB368147
Favolus roseus
PEN33 Malaysia AB735975 AB368099 AB368156
Favolus rugulosus
MP191 Brazil MN648684 MN648712
Favolus yanomami
ACM1295 Brazil MN648686 MN648714 —
Lentinus bertieri
TENN59773 Dominican
Republic GU207303 AY615984 —
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Taxon Voucher Country GenBank accession number
ITS nc LSU
rDNA
RPB2
Lentinus crinitus
DSH9243C Costa Rica KP283495 KP283523 —
Lentinus tigrinus
MUCL22821 Belgium AB478881 AB368072 AB368130
Megasporoporia
cavernulosa
Wu9508328 China — AY333800 —
Mycobonia
brunneoleuca
GAS625 Brazil MZ997324 MZ996883 OK032602
Mycobonia
brunneoleuca
TENN57579 Costa Rica AY513570 AJ487934 —
Mycobonia cucullata
MP204 Brazil MZ997329 MZ996888
Mycobonia cucullata
TENN58910 Argentina AF516600 AJ488124 —
Mycobonia ava
MP207 Brazil MZ997326 MZ996885 —
Mycobonia ava
MP213 Brazil MZ997325 MZ996884 —
Mycobonia ava
TENN59088 Argentina AY513571 AJ487933 —
Mycobonia miquelii
GAS1122 Brazil MZ997327 MZ996886 —
Mycobonia miquelii
VOG213 Brazil MZ997328 MZ996887 OK032603
Mycobonia orientalis
Cui8707 China KX880623 KX880662 —
Mycobonia orientalis
Dai13584 China KX900071 KX900185 —
Mycobonia orientalis
WD2157 Japan AB587637 AB368114 AB368170
Neodatronia
gaoligongensis
Cui8055 China JX559269 JX559286 JX559317
Neodatronia sinensis
Dai11921 China JX559272 JX559283 JX559320
Neodictyopus
atlanticae
DS1286 Brazil KY631763 KY631774 KY744950
Neodictyopus
dictyopus
GAS272 Brazil KY631766 KY631777 KY744952
Neofavolus alveolaris
WD2340 Japan AB735970 AB368077 AB368135
Neofavolus
cremealbidus
TUMH 50009 Japan AB735980 AB735957 —
Neofavolus mikawae
TFMF-27417 Japan AB735963 AB735943 —
Page 7/24
Taxon Voucher Country GenBank accession number
ITS nc LSU
rDNA
RPB2
Neofavolus
suavissimus
ADD7 USA KP283501 KP283527 —
Neofavolus
subpurpurascens
CG6242 Brazil MH544277 MH544275 —
Picipes austroandinus
MR10701 Argentina AF516569 — —
Picipes badius
WD2341 Japan AB587625 AB368083 AB368140
Picipes tubaerformis
WD1839 Japan AB587634 AB368101 AB368158
Polyporellus
arcularius
RGT830522 Canada AF516523 AB368081 AB368138
Polyporellus brumalis
TENN61760 USA FJ596883 AB368084 AB368141
Polyporellus ciliatus
TENN57698 Denmark AB070882 AJ487943 —
Polyporus tricholoma
TENN56503 Puerto Rico AF516555 AB368100 AB368157
Polyporus tuberaster
DAOM7997B USA AY218420 AF261544 —
Polyporus tuberaster
WD2382 Japan AB474086 AB368104b AB368161
Polyporus varius
WD619 Japan AB587635 AB368110 AB368167
Trametes hirsuta
RLG5133T USA JN164941 JN164801 JN164854
Trametes versicolor
FP135156 USA JN164919 JN164809 JN164850
ML analysis was performed on the CIPRES SCIENCE GATEWAY 3.1 (Miller et al. 2010) using RAxML 8.1.4
(Stamatakis 2014). The analysis rst involved 100 ML searches, each starting from one randomized
stepwise-addition parsimony tree, under a GTRGAMMA model, with all other parameters estimated by the
software. We provided a partition le with the dened partitions to force RAxML to search for a separate
evolution model for each partition. Bootstrap support values (BS) were obtained under the same models
and partitioning schemes allowing the program to automatically halt bootstrapping by using the
autoMRE option. A node was considered to be strongly supported if it showed BPP ≥ 0.95 and/or BS ≥
80%.
Trametes hirsuta
(Wulfen) Pilát and
T. versicolor
(L.) Lloyd were used as the outgroup based on
Motato-Vásquez et al. (2018).
Ancestral character state reconstruction of hymenophore
conguration
In order to reconstruct hymenophore evolution of the phylogeny of
Polyporus s.l.
we evaluated two
models of character state evolution: Equal Rates (ER) and All Rates Different (ARD) using the tDiscrete
function of the R package Geiger 2.0 (Harmon et al., 2008). We selected the best model based on the
Page 8/24
corrected Akaike Information Criterion (AIC) and used this for the reconstruction. The evolutionary
analysis of the hymenophore was performed by stochastic character mapping (Huelsenbeck et al. 2003;
Bollback 2006) implemented in the R package Phytools (Revell 2012), using the make.simmap and
describe.simmap functions (Revell 2012). A total of 1000 stochastic maps were generated. Results were
subsequently summarized and plotted on the phylogeny using the R package Ape 5.3 (Paradis et al.
2004). Hymenophore types were assigned as: circular pores = 0, angular pores = 1, subporoid lamellae = 2,
lamellate = 3, stereoid = 4 (Supp Table).
Results
Phylogenetic analyses
Phylogenetic analyses included 57 terminals representing 48 putative species. The nal alignment
consisted of 2481 characters, of which 1389 were conserved and 908 were parsimony-informative. Best
partitioning scheme was six subsets: ITS1-ITS2, 5.8S, nc LSU rDNA,
RPB2
1st codon position, 2nd codon
position and 3rd codon position. The evolutionary model selected were GTR + I + G (ITS1 and ITS2), SYM
+ I + G (5.8S), GTR + I + G (nc LSU rDNA), GTR + G (
RPB2
1st codon position), HKY + I + G (
RPB2
2nd codon
position) and GTR + I + G (
RPB2
3rd codon position). The nal alignment was deposited at TreeBASE
(28731).
Our phylogenetic analyses showed that
Pseudofavolus
and
Mycobonia
represent articial groups
according to current circumscriptions, as
Pseudofavolus
species actually nest in a clade together with
Mycobonia
species, which are phylogenetically distant from
Polyporus s.s.
(Fig. 1). Therefore, in order to
accommodate these species in a natural group based upon priority of publication, we propose to transfer
Pseudofavolus
species into the genus
Mycobonia
(see Taxonomy section). Results from phylogenetic
analyses recovered ve strongly supported linages within the
Mycobonia
clade, representing ve species
that can also be recognized by morphological and ecological differences:
M. brunneoleuca
(PP = 1.0, BS
= 99),
M. cucullata
(PP = 1.0, BS = 96),
M. ava
(PP = 1.0, BS = 91),
M. miquelii
(PP = 1.0, BS = 96) and the
new species
M. yuchengii
(PP = 1.0, BS = 100).
Mycobonia yuchengii
clade is composed by Asian
specimens previously identied as “
P. cucullatus”
, which will be treated below in the Taxonomy section.
Phylogenetic relationships between
Mycobonia
species are not well resolved and ITS, nc LSU rDNA and
RPB2
sequences are highly conserved in
Mycobonia
clade species, with only 100 variable sites of the
2481 in nal alignment. Additionally, the following were recovered as monophyletic:
Atroporus
(PP = 1.0,
BS = 100),
Bresadolia
(PP = 1.0, BS = 100),
Datronia
(PP = 1.0, BS = 97),
Datroniella
(PP = 1.0, BS = 100),
Dichomitus
(PP = 1.0, BS = 100),
Echinochaete
(PP = 1.0, BS = 100),
Favolus
(PP = 1.0, BS = 100),
Lentinus
/
Polyporellus
clade (PP = 1.0, BS = 100),
Neodictyopus
(PP = 1.0, BS = 100),
Neofavolus
(PP = 1.0,
BS = 82), and
Picipes
(PP = 1.0, BS = 100).
Ancestral character state reconstruction of hymenophore
conguration
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The best tting model for hymenophore conguration evolution was estimated to be the ER (AIC = 93.25),
compared with ARD (AIC = 107.55). In the supplemental gures, we present estimates of the
hymenophore type in
Polyporus s.l
. based on stochastic mapping of ancestral states. Probabilities are
also showed in (Supplementary Fig. 1). The stochastic mapping of this hymenophore transition
presented high probabilities of internal nodes, but low probabilities of back-bone nodes. Furthermore, the
mapping indicated that the ancestral state for
Mycobonia
clade is estimated to have had angular pores
(0.92), as well as for
Neofavolus
(1), as previously shown by Seelan et al. (2015). For the
Lentinus
/
Polyporellus
clade we found that the probability for circular pores was low, only 0.22, though in
contrast Seelan et al. (2015) did estimate them to be circular.
Polyporus s. l.
likely had angular pores
(0.72). The number of transitions from circular to angular pores was estimated to be four, while the
number from angular to circular was three, and there was a single transition from angular to stereoid
which was placed in the
Mycobonia
clade.
Taxonomy
Mycobonia
Pat., Bull. Soc. mycol. Fr. 10(2): 76 (1894), emend. Palacio & Westphalen.
=
Pseudofavolus
Pat., Essai Tax. Hyménomyc. (Lons-le-Saunier): 80 (1900)
Type species:
Mycobonia ava
(Sw.) Pat.
Basidiomata annual, solitary to imbricate, pileate, abelliform, cucullate, conchate-abelliform, subsessile
with a very short lateral stipe to sessile; exible to hard when fresh, hard and boney consistency when dry;
pilear surface glabrous, smooth to tessellate (when poroid), pale ochraceous to brownish ochraceous;
context thin up to 3 mm, homogeneous, beige. Hymenophore surface poroid or stereoid with hyphal pegs;
when poroid pores large angular to circular, 0.2–3 mm wide, dissepiments entire to lacerated; tubes short
up to 4 mm long; when stereoid with sterile prominent hyphal pegs up to 180 µm long. Stipe reduced to
subdiscoid, attached to substrate by a basal disc, concolorous with the pilear surface, to absent (Fig. 2).
Hyphal system dimitic; generative hyphae 2–5 µm wide, with clamp connections, hyaline, thin-walled,
scanty, CB−, IKI−; skeletal-binding hyphae 2–6 µm wide, densely interwoven, thick-walled, exuous,
densely branched and interwoven dominating throughout the basidiomata, CB − to CB+, IKI − to slightly
dextrinoid in mass. Pileipellis not well differentiated. Cystidia not seen. Dendrohypidia present in the
dissepiments. Basidia clavate, with four sterigmata. Basidiospores large, up to 25 × 11 µm, ellipsoid to
cylindrical, thin-walled, smooth, hyaline, IKI−, CB− (Fig. 3).
Ecology and distribution: growing on fallen branches of unidentied angiosperms, causing white-rot;
known from tropical to subtropical areas.
Comments:
Mycobonia
is characterized by the pileate, cucullate and boney consistency of the
basidiomata with the hymenophore being stereoid with hyphal pegs or poroid with shallow tubes pores
(Fig. 2), and large basidiospores (Fig. 3).
Page 10/24
Mycobonia cucullata (Mont.) Palacio & R.M. Silveira, comb. nov.
MycoBank: MB841083
≡
Favolus cucullatus
Mont., Annls Sci. Nat., Bot., sér. 2 17: 125 (1842)
≡
Pseudofavolus cucullatus
(Mont.) Pat., Essai Tax. Hyménomyc. (Lons-le-Saunier): 81 (1900)
≡
Hexagonia cucullata
(Mont.) Murrill, Bull. Torrey bot. Club 31(6): 332 (1904)
≡
Polyporus miquelii
var.
cucullatus
(Mont.) Corner, Beih. Nova Hedwigia 78: 90 (1984)
=
Favolus curtipes
Berk. & M.A. Curtis, Hooker's J. Bot. Kew Gard. Misc. 1: 234 (1849)
=
Polyporus curtipes
(Berk. & M.A. Curtis) Ryvarden, Syn. Fung. (Oslo) 5: 213 (1991)
Description: Basidiomata annual, solitary to imbricate, laterally short stipitate, dimidiate to subsessile.
Pileus 15–40 mm from the base to margin of the pileus, 30–60 mm wide, and 2 mm thick; dimidiate to
abelliform, conchate-abelliform when dry, exible when fresh, very tough when dry; surface glabrous,
smooth or slightly tessellate, not zoned, beige to pale ochraceous, slightly reddish brown towards the
margin when dry; margin entire to mbriate, involute when dry, pale ochraceous; context of the pileus up
to 2 mm wide, homogeneous, beige. Hymenophore surface poroid, light brown, pores 2–3 per mm,
angular to circular; dissepiments entire to lacerated; tubes up to 1 mm long. Stipe reduce subdiscoid up to
2 mm long, concolorous with pilear surface, or practically non-existent and attached to the substrate by a
basal disc up to 10 mm in diam.
Hyphal system dimitic; generative hyphae 2–4 µm wide, with clamp connections, hyaline, thin-walled,
scanty, CB−, IKI−; skeletal-binding hyphae 2–6 µm wide, densely interwoven, thick-walled, exuous,
branched 4–6 times, branches tapering to liform tips 0.5 µm wide, dominating throughout the
basidiomata, CB−, IKI − to slightly dextrinoid in mass in the dissepiments. Hyphal pegs absent. Pileipellis
not well differentiated. Cystidia not seen; hymenium with some nal tips of skeletal-binding hyphae
among the basidia. Basidia 25–36 × 7–14 µm, mostly clavate, with four sterigmata. Basidiospores 14–
20 × 5–7 µm (m = 17.5 × 5.7 µm) Q = 2.3–4 (m = 3; n = 100/5), narrowly cylindrical to sub-cylindrical, thin-
walled, smooth, hyaline, IKI−, CB−.
Ecology and distribution: growing on fallen branches of unidentied angiosperms; known from Argentina
(Robledo and Rajchenberg 2007 as
Polyporus curtipes
), Brazil (Baltazar and Gibertoni 2009 as
Pseudofavolus cucullatus
), Costa Rica (Velázquez and Ruíz-Boyer 2005 as
Pseudofavolus cucullatus
),
Cuba (type locality), and Mexico (Nava-Mora and Valenzuela (1997) as
Polyporus curtipes
).
Specimens examined: Brazil, Paraná, Foz do Iguaçu, Parque Nacional do Iguaçu, 25°37'21.1"S,
54°28'11.0"W, 22 Jan 2017, leg. M. Palacio 204, (ICN197323). — Cuba, leg. Ramón de la Sagra & C.
Wright (PC) (holotype of
Pseudofavolus cucullatus
).
Page 11/24
Comments:
Mycobonia cucullata
can be distinguished by the hexagonal pores 2–3 per mm, short tubes
up to 0.7 mm long, thinner context up to 0.2 mm wide, and by the narrowly cylindrical basidiospores. This
species has been recorded in South America under the name
Favolus curtipes
Berk. & M.A. Curtis (Silveira
and Wright 2005). However, according to the protologue of
F. curtipes
, this species can be differentiated
from
M. cucullata
by the smaller and less rigid pores, eshier context and darker pilear surface (Hooker
1849).
Mycobonia miquelii (Mont.) Palacio & Westphalen, comb. nov.
MycoBank: MB841084
≡
Polyporus miquelii
Mont., Annls Sci. Nat., Bot., sér. 3 4: 357 (1845)
≡
Hexagonia miquelii
(Mont.) Sacc., Syll. fung. (Abellini) 6: 361 (1888)
≡
Scenidium miquelii
(Mont.) Kuntze, Revis. gen. pl. (Leipzig) 3(3): 516 (1898)
≡
Pseudofavolus miquelii
(Mont.) Pat., Essai Tax. Hyménomyc. (Lons-le-Saunier): 81 (1900)
=
Favolus induratus
Berk., Ann. Mag. nat. Hist., Ser. 2 9: 197 (1852)
=
Hexagonia indurata
(Berk.) Murrill, Bull. Torrey bot. Club 31(6): 332 (1904)
=
Favolus daedaleoides
Speg., Revista Argent. Hist. Nat. 1(2): 108 (1891)
Description: Basidiomata annual, solitary, laterally short stipitate to subsessile. Pileus 20–40 mm from
the base to margin of the pileus, 20–60 mm wide, and 2–4 mm thick; exible when fresh, very tough and
convex when dry, abelliform, conchate-abelliform; surface glabrous, tessellate bullate-reticulate, not
zoned, pale ochraceous to brownish ochraceous with reddish-brown spots mainly towards the margin;
margin entire to lacerate, wavy, incurved when dry, reddish-brown; context of the pileus 2–3 mm wide,
homogeneous, beige. Hymenophore surface poroid, pale ochraceous to brownish ochraceous, pores 1–
1.5 per mm, circular to angular; dissepiments entire to lacerated; tubes up to 4 mm long. Stipe reduce
subdiscoid up to 2 mm long, concolorous with pilear surface, or practically non-existant and attached to
substrate by a basal disc up to 7 mm in diam.
Hyphal system dimitic; generative hyphae 2–5 µm wide, with clamp connections, hyaline, thin-walled,
scanty, CB−, IKI−; skeletal-binding hyphae 2–6 µm wide, densely interwoven, thick-walled, exuous,
branched 4–7 times, branches tapering to liform tips − 1 µm wide, dominating throughout the
basidiomata, CB−, IKI−. Hyphal pegs absent. Pileipellis not well differentiated. Cystidia not seen;
hymenium with some nal tips of skeleto-binding hyphae between the basidia. Basidia 20–35 × 9–15
µm, mostly clavate, with four sterigmata. Basidiospores 11–17 × 5.5–8.5 µm (m = 14 × 6.5 µm) Q = 2.3–
2.8 (m = 2.4; n = 100/5), sub-cylindrical, thin-walled, smooth, hyaline, IKI−, CB−.
Page 12/24
Ecology and distribution: growing on fallen branches of unidentied angiosperms; common species
originally described from Surinam and also registered from Argentina, Brazil (Capelari and Maziero 1988),
Costa Rica (Velázquez and Ruíz-Boyer 2005), and the Dominican Republic.
Specimens examined: Brazil, Espírito Santo, Santa Tereza, Rebio Augusto Ruschi, Trilha da Cachoeira, 13
Dez 2016, leg. A. Magnago 1305 (ICN197324); Rio Grande do Sul, Viamão, Parque Estadual de Itapuã, 21
Apr 2017, leg. G. Alves-Silva 1120 (ICN197320); ibid, leg. G. Alves-Silva 1121 (ICN197321); ibid, leg. G.
Alves-Silva 1122, (ICN197322); Guaiba, Fazenda Maximiniano, 11 Nov 2017, leg. V. Oliveira-Garcia 213
(ICN197423); Porto Alegre, Morro Santana, 14 Dez 2007, leg. M. Westphalen 85/07 (ICN134137); Santa
Catarina, Florianópolis, Parque Ecológico Córrego Grande, 27°35'55''S, 48°30'36''W, 10 May 2018, leg. M.
Monteiro 160 (FLOR); São Paulo, Cananéia, Parque Estadual da Ilha do Cardoso, Trilha do Poço da Anta,
7 Jan 2019, leg. M.P. Drewinski 463, (SP499085); ibid., Parque Estadual das fontes do Ipiranga,
23°38'36.1"S, 46°37'21.6"W, 28 Mar 2019, leg. M.P. Drewinski 459 (SP499083).
Comments: Large and angular pores, 1–1.5 per mm, and a tessellate pilear surface with a reddish-brown
margin, which differentiates
M. miquelii
from other
Mycobonia.
Mycobonia yuchengii Yuan Yuan & Palacio, sp. nov. Figure 4.
MycoBank: MB841082
Type. — China, Hainan Province, Lingshui County, Diaoluoshan Forest Park, on fallen angiosperm trunk,
13 Jun 2014, leg. Dai 13584A (BJFC017323).
Etymology: — Yuchengii (Lat.): in honor of the Chinese mycologist, Prof. Yu Cheng Dai.
Description: Basidiomata annual, pileate, attached to substrate with a short laterally stipe-like base,
leathery when fresh, becoming hard or rigid upon drying. Pileus dimidiate to abelliform, projecting up to
3.5 cm, 4 cm wide, 1.5 mm thick at the center. Pilear surface cream to pale yellow-brown, glabrous,
azonate, sometimes radiate-striate; margin acute, wavy when dry. Pore surface yellow-brown to umber;
pores irregular, sometimes hexagonal, 1–3 per mm; dissepiments thin, entire to slightly lacerate. Context
pale yellow-brown, hard corky, thin, less than 0.5 mm thick. Tube layer concolorous with the poroid
surface, up to 2 mm long. Stipe short, concolorous with the pileal surface, up to 0.5 cm long, often
forming a small disc at the base.
Hyphal system dimitic; generative hyphae bearing clamp connections; skeletal-binding hyphae dominant,
thick-walled, frequently branched, slightly dextrinoid, especially at dissepimental edge, CB+, tissue
unchanged in KOH. Contextual generative hyphae infrequent, hyaline, thin-walled, rarely branched, 2.5–
3.5 µm in diam; contextual skeletal-binding hyphae dominant, hyaline, thick-walled with a wide to narrow
lumen, some of them sub-solid, exuous, frequently branched, tightly interwoven, 2.5–4 µm in diam.
Tramal generative hyphae scanty, hyaline, thin-walled, rarely branched, 2–3.5 µm in diam,; tramal skeletal-
binding hyphae thick-walled to almost solid, frequently branched, tightly interwoven, 2–4.5 µm in diam.
Cystidia and cystidioles absent. Dendrohyphidia present along the hymenium, especially abundant in the
Page 13/24
dissepiments. Basidia broadly clavate, with four sterigmata and a basal clamp connection, 20–35 × 12–
15 µm; basidioles in shape similar to basidia, but slightly smaller. Basidiospores ellipsoid to mango-
shaped, hyaline, thin-walled, smooth, sometimes with one or two guttule, IKI–, CB–, (11)13–17(18) ×
(5.5)5.8–8.5(8.8) µm, L = 14.8 µm, W = 7.1 µm, Q = 2.08–2.25 (n = 60/2).
Ecology and distribution: When Wu et al. (2020) made a systematic study on the polypores from
subtropical and tropical China,
M. yuchengii
was found as a common species in South China, usually
grows on angiosperm trunks or branches and occurring in open areas of the forest.
Specimens examined: China, Hainan Province, Wuzhishan County, Wuzhishan Forest
Park, on fallen angiosperm trunk, 10 Jun 2016 Dai 16493 (BJFC022610); Hunan
Province, Wulingyuan District, on fallen angiosperm trunk, 17 Ago 2010, leg. Dai 11682
(BJFC008806); Guangdong Province, Shixing County, Chebaling Nature Reserve, on fallen angiosperm
branch, 23 Jun 2010, Cui 8707 (BJFC007647); ibid., 23 Jun 2010, leg. Cui 8747 (BJFC007687); Yunnan
Province, Mengla County, Xishuangbanna Botanical Garden, on fallen angiosperm branch, 23 Jul 2014,
leg. Dai 13893 (BJFC017623); ibid., leg. Dai 13894 (BJFC017624); Green Stone Forest Park, on fallen
angiosperm branch, 4 Ago 2005, leg. Dai 6678 (IFP015327); Puer County, Laiyanghe Forest Park, on
fallen angiosperm trunk, 6 Jun 2011, leg. Dai 12207 (BJFC010490).
Comments:
Mycobonia yuchengii
was previously identied as “
P. cucullatus
” (Dai 2007).
Mycobonia
yuchengii
basidiomata have shallow pores, so macroscopically it resembles species of
Grammothele
Berk. & M.A. Curtis. In addition, species in both genera have dendrohyphidia along the hymenia, and their
skeletal hyphae are dextrinoid (Zhou and Dai 2012). However,
Grammothele
lacks both stipe and binding
hyphae.
Mycobonia brunneoleuca (Berk. & M.A. Curtis) Pat., Enum. Champ. Guadeloupe (Lons-le-Saunier): 23
(1903).
≡
Hydnum brunneoleucum
Berk. & M.A. Curtis, Trans. Linn. Soc. London
22: 129. 1857, non
Polyporus brunneoleucus
Berk. 1846.
≡
Polyporus polyacanthophorus
Nakasone, Mycotaxon 130(2): 383. 2015.
Description: see Nakasone (2015).
Ecology and distribution: growing on fallen branches of unidentied angiosperms, in cloud forest at 700–
2700 m.a.s.l. in Brazil (Gerlach and Loguercio-Leite 2011), Colombia, Costa Rica, Honduras (Nakasone
2015), Martinique (Burt 1919), Panamá (Martin 1939), Paraguay, Puerto Rico (Nakasone 2015), and
Venezuela (type locality).
Page 14/24
Specimens examined: Brazil, Rio Grande do Sul, São Francisco de Paula, Floresta Nacional de São
Francisco de Paula, 25 May 2019, leg. M. Palacio 438 (ICN202959); Santa Catarina, Joaçaba, Parque
Ecológico Municipal Rio do Peixe, leg. G. Alves-Silva 625, (FLOR60335).
Comments:
Mycobonia brunneoleuca
is principally characterized by the large basidiospores 15–24(25) ×
(6)7–11 µm, narrowly ellipsoid, thin walled and hyaline. Macroscopically is characterized by the concave
and shell-like basidiomata with stereoid hymenophore projecting sterile hyphal pegs, which is almost
identical to
M. ava
.
Mycobonia brunneoleuca
and
M. ava;
can be differentiated by the basidiospores,
which are sub-cylindrical and smaller (up to 5–7 µm wide) in
M. ava
, and also by the ecological niches
that these two species occupy (Gerlach and Loguercio-Leite 2011).
Mycobonia brunneoleuca
occurs in
cloud forests of the Caribbean, and throughout Central and South America between 700 and 2,700 m asl.,
while
M. ava
occurs at elevations up to 700 m asl. from North America to Argentina. In addition to these
morphological and ecological differences, our phylogenetic analysis showed that
M. brunneoleuca
and
M. ava
represent two different linages (Fig. 1).
Mycobonia ava (Sw.) Pat., Bull. Soc. mycol. Fr. 10(2): 77. 1894.
≡
Peziza ava
Sw.: Fr., Prod.: 150. 1788.
≡
Hydnum avum
(Sw.: Fr.) Berk., Ann. Mag. Nat. Hist., 10: 380. 1843 [“1842”].
≡
Bonia ava
(Sw.: Fr.) Henn., Hedwigia 36: 192. 1897.
≡
Auricularia ava
(Sw.: Fr.) Farl., Bibl. Index N. Amer. Fung.: 307. 1905.
≡
Grandinioides ava
(Sw.: Fr.) Banker, Mem. Torrey Bot. Club 12: 179. 1906.
≡
Polyporus curtipes
subsp.
avus
(Sw.: Fr.) D. Krüger, Cryptog. Mycol. 31: 399. 2010.
≡
Polyporus epitheloides
Nakasone, Mycotaxon 130(2): 383. 2015.
Description: see Julich (1976).
Ecology and distribution: growing on fallen branches of unidentied angiosperms, in Argentina, Colombia
(Jülich 1975), Cuba (Burt 1919), Jamaica (Swartz 1788) (type locality), North America (Burt 1919), Brazil
(Jülich 1975; Baltazar and Gibertoni 2009) and Venezuela (Corner 1984).
Specimens examined: Brazil, Amazonas, Novo Aripuanã, BR-230, 23 Apr 1985, leg. K.F. Rodrigues 325
(INPA136956); Paraná, Foz do Iguaçu, Parque Nacional do Iguaçu, 25°37'21.1"S, 54°28'11.0"W, 23 Jan
2017, leg. M. Palacio 207 (ICN197318); ibid., leg. M. Palacio 213, (ICN197319); Rio Do Janeiro, Rio de
Janeiro, Parque Nacional da Tijuca, 8 Feb 2017, leg. F.T.F. Linhares 228 (ICN197346); São Paulo, São
Paulo, Parque Estadual da Cantareira, Trilha da cachoeira, 25 Mar 2019, leg. M.P. Drewinski 449,
(SP499082).
Page 15/24
Comments: Microscopically
M. ava
is very similar to
M. miquelii
, especially by the sub-cylindrical
basidiospores, however
M. ava
can be differentiated by the hymenophore being stereoid with hyphal
pegs, while in
M. miquelii
it is poroid.
Other species possibly included in the genus Mycobonia:
Favolus curtipes
Berk. & M.A. Curtis, Hooker's J. Bot. Kew Gard. Misc. 1: 234 (1849)
Favolus tenuis
Fr., Syst. orb. veg. (Lundae) 1: 76 (1825)
Hexagonia bipindiensis
Henn., Bot. Jb. 38(1): 122 (1905) [1907]
Hexagonia pulchella
Lév., Annls Sci. Nat., Bot., sér. 3 2: 200 (1844)
Polyporus orinocensis
Pat. & Gaillard, Bull. Soc. mycol. Fr. 4(2): 31 (1888)
Polyporus polygrammus
Mont., Annls Sci. Nat., Bot., sér. 2 8: 365 (1837)
Pseudofavolus nigrus
Ryvarden, Mycotaxon 28(2): 537 (1987)
KEY TO MYCOBONIA NEOTROPICAL SPECIES
1 Hymenophoral surface poroid 2
1* Hymenophoral surface stereoid with hyphal pegs 5
2 Pores 1–1.5 per mm; pilear surface tessellate, bullate-reticulate, pale ochraceous to brownish
ochraceous with reddish-brown spots mainly towards the margin ...........................
M. miquelii
2* Pores 2–3 per mm; pilear surface smooth to slightly tessellate, or radially wrinkled, beige to pale
ochraceous, to chesnut to black 3
3 Pores 0.2–0.5 per mm; pilear surface smooth or slightly tessellate, beige to pale ochraceous, slightly
reddish brown towards the margin when dry; known from Mexico to Argentina
M. cucullata
3* Pores 4–5 per mm; pilear surface smooth darker, only known from Venezuela. 4
4 Pilear surface pale chestnut to reddish-brown
M. yuchenguii
4* Pilear surface black
P. nigrus
5 Basidiospores narrowly ellipsoid 15–24(25) × (6)7–11 µm; growing in cloud forests of the Caribbean
and Central and South America between 700 and 2,700 m asl.
M. brunneoleuc
a
Page 16/24
5* Basidiospores sub-cylindrical 12–18 × 5–7 µm; growing at elevations up to 700 m asl. from North
America to Argentina
M. ava
Discussion
Mycobonia
and
Pseudofavolus
in their traditional circumscriptions are paraphyletic groups. The
differences between ITS, nc LSU rDNA and
RPB2
sequences within
Mycobonia
species are signicantly
small, with just 100 variable sites from 2481 characters, thus showing that the phylogenetic separation
between both genera is articial. Following the guidelines to dene fungi genera formulated by Vellinga et
al. (2015), we consider
Pseudofavolus
a synonym of
Mycobonia
(older name with priority). Therefore, we
emended
Mycobonia
to encompass stereoid and also poroid species. We combined
M. cucullata
and
M.
miquelii
since we could assess phyllogenetic data of those species through molecular analysis.
Regarding other
Pseudofavolus
species, such as
P. nigrus
and
P. orinocensis
, we preferred to avoid
nomenclatural changes until sequences of those species become available and their phylogenetic
position can be conrmed, especially in a group with several genera with similar morphology.
Phylogenetic relationships within
Mycobonia
clade are still not well resolved, especially due to the
similarity in the sequences among the different species. More molecular data including other DNA
regions could help clarify that in the future. However,
Mycobonia
species can be differentiated and
recognized by ecologically through distribution patterns and elevation, while morphologically through the
basidiospore shape and size, as well as the hymenophore type.
Hymenophoral transitions within genera have often been documented in other groups of Polyporales, as
Antrodia
(Runnel et al. 2019),
Steccherinum
(Westphalen et al. 2018; Miettinen et al. 2012),
Metuloidea
(Miettinen and Ryvarden 2016), and also within the core polyporoid clade in
Neofavolus
,
Lentinus
/
Polyporellus
(Seelan et al. 2015). Additionally, Seelan et al. (2015) estimated the ancestral
hymenophoral conguration for
Neofavolus
and
Lentinus
/
Polyporellus
. However, ancestral character
state reconstruction including both stereoid and poroid hymenophore had not been estimated previously
in the core polyporoid clade. Our results suggest a transition from angular pores to stereoid hymenophore
in
Mycobonia
clade, and that the plesiomorphic condition for
Polyporus s.l.
is to have angular pores.
Other groups in the core polyporoid clade can also present stereoid hymenophore with hyphal pegs, as
Dichomitus
,
Ephitele
, and
Gramothele
(Nakasone 2015). However, these groups present resupinate
basidiomata, while the basidiomata in
Mycobonia
are pileate. Specically, hyphal pegs are also present
in other clades of Polyporaceae, as
Favolus
,
Lentinus
, and
Neofavolus
, which are different in presenting
basidiomata with lamellate hymenophore. The presence of hyphal pegs could represent an anatomical
advantage, considering it has been suggested that there exists a possible relation with moisture retention
and also protection from insect invasion (Pegler 1983). However, the function of the hyphal pegs is still
unknown (Pegler and Young 1983), and it is not a homologous character (Hibbett and Vilgalys 1993).
Mycobonia
shares with
Polyporus s.l.
a dimitic hyphal system with skeletal-binding hyphae, hyaline, and
thin-walled basidiospores, and caising white-rot, which appear to be a conserved characteristic across all
Page 17/24
Polyporus s.l.
species. However, relationships between genera, like
Mycobonia
in
Polyporus s.l.
are yet to
be solved.
Declarations
ACKNOWLEDGEMENTS
The authors would like to thank the Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio)
for giving permission to sample the collections and also the curators of the herbaria mentioned above for
the specimen loans. We would like to thank Felipe Bittencourt and Genivaldo Alves Silva for sharing
photos and Hunter Daniel for for the English revision of the manuscript. We also thank to Professor Yu-
Cheng Dai and Xiao-Hong Ji for their assistance and support in the molecular procedures performed at
the Institute of Microbiology (Beijing Forestry University, China). Rosa Mara Borges da Silveira is
supported by Conselho Nacional de Desenvolvimento Cientíco e Tecnológico (process no.
308122/2019-4). Melissa Palacio is grateful for Coordenacão de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES), which provided a PhD scholarship, as well as the International Association for Plant
Taxonomy for an IAPT research grant.
Funding: Coordenacão de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) provided a PhD
scholarship to Melissa Palacio, and the International Association for Plant Taxonomy awarded her an
IAPT Research Grant.
Conicts of interest/Competing interests: Not applicable
Availability of data and material: the manuscript data are deposited in world-reference repositories
(GenBank, TreeBase) and the fungal collections are deposited in herbaria recognized by the Index
Herberiorum.
Code availability: Not applicable
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Figures
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Figure 1
Phylogenetic relationships in Polyporus s.l. from ITS, nc LSU rDNA and RPB2 sequences. Topology from
ML analysis. Bayesian posterior probability above 0.95 and bootstrap values above 80% are shown. Pies
at nodes represent probability of each state from the summary of stochastic mapping of ancestral state
estimates of hymenophore conguration across Polyporus s.l. (blue = circular pores, green = angular
pores, orange = subporoid lamellae, pink = lamellae, grey = stereoid).
Figure 2
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Basidiomata and hymenophore detail of Mycobonia species. A–D. Mycobonia brunneoleuca (GAS625).
E. Mycobonia ava (MP207). F–H. M. miquelii (Monteiro 160; GAS 1122). I–J. M. cucullata. K. M.
yuchengii (type Dai 13584A). Photo A. by G. Alves-Silva and F. by Felipe Bittencourt.
Figure 3
Microscopic features of Mycobonia species. A. Hymenium, tramal hyphae and sterile hyphal pegs in M.
ava (MP207). B–E. Basidiospores. B. M. ava. C. M. brunneoleuca. D. M. cucullata. E. M. miquelii.
Page 23/24
Figure 4
Microscopic features of Mycobonia yuchengii from type Dai 13584A (BJFC017323). A. Basidiospores. B.
Basidia. C. Dendrohypidia. D. Hyphae from the trama. E. Hyphae from the context.
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