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© 2018 Westerdijk Fungal Biodiversity Instute 341
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal Biodiversity Institute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
Fungal Systemacs and Evoluon
doi.org/10.3114/fuse.2018.02.10
VOLUME 2
DECEMBER 2018
PAGES 341–359
INTRODUCTION
Phylloporus is a genus of lamellate fungi in the family Boletaceae
that is primarily distributed throughout the tropics (Kirk et
al. 2008, Neves & Halling 2010, Neves et al. 2012, Zeng et al.
2013). Singer (1945b) moved Phylloporus to the subfamily
Xerocomoideae due to its Phylloporus-type hymenophoral
trama and olive-brown spore print. Bresinsky & Besl (2003)
synonymised Phylloporus with Xerocomus based on molecular
data, though only a few taxa were sampled in that study. Recent
support the monophyly of Phylloporus, showing that Xerocomus
et al. 2013, Wu et al.
of Phylloporus in an expanded Xerocomoideae that now also
includes taxa with Boletus-type hymenophoral trama and pale
et al.
Phylloporus to
include about 70 species, but several subsequent studies have
et al. 2010, 2012,
García-Jiménez 2013, Zeng et al. 2013, Ye et al. 2014, Hosen & Li
2015, 2017). Species of Phylloporus form ectomycorrhizal (ECM)
Casuarinaceae, Dipterocarpaceae,
Fabaceae, Fagaceae, Myrtaceae, and Pinaceae (Heinemann
Watling 2008, Neves & Halling 2010, Neves et al. 2012, García-
Jiménez 2013, Zeng et al. 2013, Ye et al. 2014, Hosen & Li 2015,
2017). Five species of Phylloporus have been described from
North America (Neves 2007, Neves et al. 2010).
Phylloporus bolenoides is a lamellate to subporoid bolete
that was formerly described based on material collected by Harry
eastern and south-eastern USA south to the Gulf Coast (Singer et
al.et al. 2000,
et
al.
Santana et al. (2007) as occurring in the Dominican Republic. It
Pinus
spp., Pinaceae), but is also found in mixed pine and oak (Quercus
spp., Fagaceae et al. 1990,
et al.
Phylloporus and Phylloboletellus are no longer alone: Phylloporopsis gen. nov. (Boletaceae),
a new smooth-spored lamellate genus to accommodate the American species Phylloporus
bolenoides
A. Farid1*§, M. Gelardi2*, C. Angelini3,4, A.R. Franck5, F. Costanzo2, L. Kaminsky, E. Ercole7, T.J. Baroni8, A.L. White1, J.R. Garey1, M.E.
Smith, A. Vizzini7§
1
2
3Via Cappuccini 78/8, I-33170 Pordenone, Italy
4
5
7
8Department of Biological Sciences, State University of New York – College at Cortland, Cortland, NY 1304, USA
*Authors contributed equally to this manuscript
§Corresponding authors: alfredo.vizzini@unito.it, arian@mail.usf.edu
Abstract: The monotypic genus Phylloporopsis is described as new to science based on Phylloporus bolenoides. This
TEF1-α, and RPB1). A
P. bolenoides from the Dominican Republic and Florida (USA) is provided along
microscopic images of anatomical features and scanning electron microscope images of basidiospores. The taxonomic
P. bolenoides
Phylloporus, Phylloboletellus,
Phyllobolites and Bothia are discussed.
Key words:
Boletales
lamellate boletes
molecular phylogeny
taxonomy
Xerocomoideae
Published online: 23 October 2018.
© 2018 Westerdijk Fungal Biodiversity Instute
Farid et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
342
Singer et al.
the species in Phylloporus sect. Fibula. The species was originally
assumed to represent an intermediate taxon between Suillus and
Bolenuset al.
placed P. bolenoides in Phylloporus sect. Manausenses but later
within the genus because of its peculiar morphological features
with special reference to the pallid-coloured hymenophore and
P. bolenoides from Belize, the
Dominican Republic and Florida were carefully examined
using morphological and molecular approaches. Outcomes
of Phylloporus
sequences from four gene regions (ITS, 28S, TEF1-α, and RPB1)
Phylloporopsis as a unique and as yet monotypic generic lineage
in the Boletaceae
genus Bothia and the sequestrate genus Solioccasus.
This study also widens the geographical extension of P.
bolenoides to the Dominican Republic, where it is reported for
Pinus occidentalis,
on decayed wood or even on trunks of living trees, determining
MATERIALS AND METHODS
Collecon site and sampling
Specimens from Belize, Dominican Republic, and the USA were
examined from several public herbaria (CFMR, F, FLAS, JBSD,
MICH, USF), as well as private herbaria [personal herbaria of
of the public herbaria follow Thiers (2018), Herbarium
Fungorum, Authors of Fungal Names (www.indexfungorum.org/
Morphological studies
4OH,
4·7H2
Microscopic anatomical features were observed and recorded
with Congo Red. All anatomical structures were measured from
were made at 1000 × using an ocular micrometer. Basidiospores
were measured directly from the hymenium of mature basidiomata,
2
part, and the length was measured from the apex (sterigmata
Line drawings of microstructures were traced free hand based on
digital photomicrographs of rehydrated material. Scanning electron
micrographs have been obtained using a JEOL JSM IT300LV (High
thickness of 400 Å.
DNA extracon, PCR amplicaon and DNA sequencing
et al. (2015). Primers ITS1F and
ITS4 (White et al. 1990, Gardes & Bruns 1993) were used for
the ITS region; primers LR0R and LR5 (Vilgalys & Hester 1990,
Rehner & Samuels 1994) were used for the 28S rDNA, EF1-983F
TEF1-α
directed RNA polymerase II subunit 1 region (RPB1) were
et al.
RPB1 Bothia clade were developed
(Table 2). A touchdown PCR was used to amplify the RPB1
region with the newly developed primer pairs RPB1-32-F/RPB1-
835-R, RPB1-147-F/RPB1-1091-R. The cycle parameters were as
Inc. (Seoul, Republic of Korea).
Sequence alignment, data set assembly and phylogenec
analyses
The sequences obtained in this study were checked and
assembled using Geneious v. 11.1.4 (Kearse et al. 2012) and
compared to those available in GenBank database by using the
Blastn algorithm (Altschul et al. 1990). Chromatograms were
examined and manually edited for accuracy. Sequences were
and accession numbers are reported in Table 1. Homologous
sequences from vouchered specimens and from environmental
samples were selected and retrieved from Halling et al. (2007),
Trappe et al. (2013), Zeng et al. (2013), Zhu et al. (2015) and
Orihara & Smith (2017). A general combined Maximum
likelihood tree including all the Boletaceae sequences deposited
in the major clades of Boletaceae as circumscribed by Wu et al.
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
343
Table 1. TEF1-α RPB1
combined datasets). Sequences newly generated for this study are highlighted in bold.
Tax on Voucher No. Locality ITS 28S TEF1-α RPB1
Afroboletus luteolus Africa - KF030238 KF030397 -
Afrocastellanoa ivoryana Mukuvisi, Zimbabwe - -
Australopilus palumanus Queensland, Australia - -
Austroboletus fusisporus HKAS75207 China - -
Austroboletus gracilis MA, USA - KF030425 KF030358
Austroboletus mutabilis BRI AQ0795793 Queensland, Australia - - KP242078
Austroboletus subvirens BRI AQ0794171 Queensland, Australia - KP242227 - KP242045
Baorangia bicolor MB07-001 NY, USA - - KF030370
Baorangia pseudocalopus China - KY418895 -
HKAS 75739 China - KJ184558 -
HKAS 75081 Nanhua County, Yunnan Prov.,
China
- -
Boletus aereus REH 8721 Humboldt County, California,
USA
- KF030339 - KF030377
Boletus edulis Be3 Bavaria, Germany - KF030282 -
BD380 Colorado, USA - - -
Boletus pulchriceps DS4514 Chiricahua Mnts, AZ, USA - KF030409 -
Boletus rufomaculatus 4414 Chestnut Ridge Park, NY, USA - KF030248
Boletus semigastroideus Auckland, New Zealand - KF030352 KF030430 -
Boletus variipes va r. fagicola A.H. Smith 4249 Cheboygan Co, Michigan, USA - JQ327014 JQ327017 -
Borofutus dhakanus HKAS 73792 Bangladesh - JQ928575 -
Bothia castanella MB 03-053 MA, USA KF030421 KF030382
NY28003 NY, USA - - -
NY, USA - - -
NY28002 NY, USA - - -
MA, USA - - -
Bothia fujianensis Fujian Prov., China - - -
Fujian Prov., China -
Buchwaldoboletus lignicola Yichun, Heilongjiang Prov.,
China
- KF112350 KF112277
Butyriboletus appendiculatus Bap1 Bavaria, Germany - JQ327025 -
Butyriboletus roseoavus HKAS 54099 China - - KF739741
Caloboletus rmus Chestnut Ridge Park, NY, USA - KF030278 KF030408
Caloboletus inedulis Erie Co., NY, USA - JQ327013 JQ327020
Chalciporus piperatus MB 04-001 Rutland State Park, MA, USA - GU187453
Fistulinella prunicolor REH 9502 Fraser Island, Queensland,
Australia
- -
Gymnogaster boletoides REH 9455 Cooloola, Queensland,
Australia
- -
Gyrodon lividus Gl1 Bavaria, Germany - AF098378 GU187701 -
Harrya chromapes HKAS 50527 Dêqên, Yunnan Prov., China - KF112437 KF112270 -
Heliogaster columellifer KPM-NC 23012 Odawara, Kanagawa Pref.,
Japan
- -
Hourangia cheoi China - KF112385 -
Hourangia microcarpa
(Wu1324)
China - -
Hourangia nigropunctata China - KF112388 KF112287 -
Hourangia sp. China - KF112453 KF112301 -
Imleria badia xb2 Bavaria, Germany - KF030357 KF030422 -
© 2018 Westerdijk Fungal Biodiversity Instute
Farid et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
344
Table 1.
Tax on Voucher No. Locality ITS 28S TEF1-α RPB1
Sweden - -
Imleria obscurebrunnea HKAS 52557 - KF112374 KF112190 KC215225
Lanmaoa angusspora HKAS 74759 China - -
Lanmaoa asiaca Heqing, Yunnan Prov., China -
Lanmaoa carminipes Erie Co., NY, USA - JQ327001 JQ327022 -
Leccinellum griseum KPM-NC 17831 Hyogo Pref., Japan - JN378508 JN378449 -
Leccinellum cremeum China - - -
Leccinellum crocipodium KPM-NC 18041 - KC552053 KC552094 -
Leccinum moncola China - KF112443 - KF112592
Leccinum scabrum KPM-NC 17840 - JN378515 JN378455 -
Leccinum versipelle KPM-NC 17833 Scotland, UK - JN378514 JN378454 -
Neoboletus magnicus HKAS 74939 Baoshan, Yunnan Prov., China - KF112320 KF112148 -
Nigroboletus roseonigrescens GDGM 43238 Guangdong Prov., China - KT220588 KT220595 -
Octaviania decimae Mt. Hiei, Kyoto Pref., Japan - JN378409 -
Octaviania kobayasii KPM-NC 17785 Mt. Kasuga, Nara Pref., Japan - JN378478 JN378420 -
Octaviania nonae KPM-NC 17748 Amami-oshima, Kagoshima
Pref., Japan
- JN378459 JN378403 -
Octaviania tasmanica Tasmania, Australia - JN378495 -
Octaviania yaeyamaensis KPM-NC 17819 Ishigaki Isl., Okinawa Pref.,
Japan
- JN378491 JN378432 -
Paragyrodon sphaerosporus Iowa, USA - GU187593 - -
Paxillus vernalis China - - -
Phylloboletellus chloephorus Veracruz, Municipio Coatepec,
El Grande, Mexico
- - -
Phylloporopsis bolenoides JBSD127411 Jarabacoa, Dominican
Republic
MH571675 MH571711 MH588312 -
JBSD127412 Jarabacoa, Dominican
Republic
MH571676 MH571712 MH588313 -
JBSD127413 Jarabacoa, Dominican
Republic
MH571677 MH571713 MH588314 -
JBSD127414 Jarabacoa, Dominican
Republic
MH571678 MH571714 MH588315 -
FLAS-F-60407 Putnam County, Florida, USA MG845193 - - -
FLAS-F-60413 Putnam County, Florida, USA MG845194 - - -
FLAS-F-61158 Putnam County, Florida, USA MH211774 - - -
Farid 617 (USF
296126)
Tampa, Florida, USA MG817716 MG817715 - MG820263
CORT014483 Mountain Pine Ridge, Belize MH571679 MH571715 MH588316 -
CORT010991 Kountze, Texas, USA - MH571716 MH588317 -
F1118420 Sarasota, Florida, USA MH571680 MH571717 - -
Phylloporus aenuatus
(holotype)
Tangail, Bangladesh - KR094791 -
Phylloporus bellus Yunnan, SW China - -
Phylloporus brunneiceps Yunnan, SW China - -
Phylloporus catenulatus Bangladesh - KR094779 KR094789 -
Phylloporus gajari Gazipur, Bangladesh - -
Phylloporus imbricatus Yunnan, SW China - -
China - KF112398 KF112299 -
Phylloporus leucomycelinus eastern USA - -
Phylloporus luxiensis Yunnan, SW China - -
HKAS 75077 China - KF112490 KF112298
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
345
Table 1.
Tax on Voucher No. Locality ITS 28S TEF1-α RPB1
Phylloporus maculatus Yunnan, SW China - -
Phylloporus pachycysdiatus HKAS 54540 Yunnan, SW China - -
Phylloporus parvisporus Yunnan SW China - -
Phylloporus pelleeri Pp1 Austria - KF030390
Phylloporus pelleeri Q7199c Slovakia - - -
K 128205 England, UK - - -
Phylloporus rhodoxanthus SAR 89.457 eastern USA - U11925 - -
MAN075 eastern USA - - -
REH8714 eastern USA - - -
MAN099 eastern USA - - -
JLM1808 eastern USA - - -
BD374 - -
Phylloporus rubeolus HKAS 52573 Yunnan, SW China - -
Phylloporus rubrosquamosus HKAS 54542 Yunnan, SW China - -
HKAS 52552 China - KF112391 KF112289 -
Phylloporus rufescens HKAS 59722 Hainan, southern China - -
Phylloporus yunnanensis HKAS 52225 Yunnan, SW China - -
Phylloporus sp. Hunan, central China - -
Fujian, SE China - -
Hainan, southern China - -
Yunnan, SW China - -
Yunnan, SW China - -
Fujian, SE China - -
Yunnan, SW China - -
Yunnan, SW China - -
Yunnan, SW China - -
Hainan, southern China - -
Porphyrellus brunneus REH 9527 Fraser Island, QLD, Australia - -
Porphyrellus porphyrosporus Mt. Tarumae, Hokkaido, Japan -
KPM-NC 25017 Rishiri Island, Hokkaido, Japan -
MB 97-023 Walhalla, Bavaria, Germany GU187734 -
Yanbian, Jilin Prov., China - KF112482 KF112243 -
Porphyrellus sp. Sanming, Fujian Prov., China - KF112480 KF112241 -
HKAS 75078 Chuxiong, Yunnan Prov., China - KF112481 KF112242 -
Pseudoboletus parasicus Bavaria, Germany - KF030443 -
Reboletus fuscus Yunnan Prov., China - JQ928580 -
Reboletus griseus Both sn NY, USA - KF030308 KF030414 KF030373
Rossbeevera griseoveluna Hyogo, Japan - KC552031 -
Rossbeevera viaspora MEL2128491 NSW, Australia - -
Royoungia boletoides NSW, Australia - -
Rubroboletus rhodosanguineus 4252 Chestnut Ridge Park, NY, USA - KF030252 KF030412 -
Solioccasus polychromus J. Trappe 15399 Australia - - -
REH 9417 Queensland, Australia - -
Spongiforma thailandica DED 7873 Thailand - -
Strobilomyces strobilaceus (as
S. occopus)
Sf1 Bavaria, Germany - JQ327037 -
Suillellus amygdalinus Mendocino Co., CA, USA - JQ327024 -
Sutorius eximius Kunming, Yunnan Prov. CHINA - KF112399 KF112207 -
Sutorius eximius REH 8594 Jardin de Dota, Costa Rica - JQ327008 JQ327027 -
© 2018 Westerdijk Fungal Biodiversity Instute
Farid et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
were restricted to the major clade including P. bolenoides
sequences and to selected genera in the Boletaceae.
TEF1-α dataset and a 28S/RPB1
P. bolenoides
in the Boletaceae. According to the results by Orihara & Smith
(2017), species of Paxillaceae were chosen as outgroup taxa for
only on an ITS dataset was restricted to the taxa closely related
to P. bolenoides; Tylopilus felleus was used as outgroup taxon
Alignments were generated for the ITS, 28S, TEF1-α, and RPB1
datasets with MAFFT (Katoh et al.
manually adjusted using Geneious v. 11.1.4 (Kearse et al. 2012). We
(Darriba et al.
inference (BI) and Maximum likelihood (ML) criteria. The BI was
et al. 2012) with
one cold and three incrementally heated simultaneous Monte
performed independently. Trees were sampled every 1 000
remaining trees of the two independent runs, a majority rule
ML analysis was performed using RAxML v. 7.3.2 (Stamatakis
GTRGAMMA algorithm. Support values from bootstrapping runs
rapid bootstrapping algorithm. BI and ML analyses were run on
the CIPRES Science Gateway (Miller et al. 2010). Only BPP values
Table 1.
Tax on Voucher No. Locality ITS 28S TEF1-α RPB1
Turmalinea persicina KPM-NC 18001 Iwakura Kyoto Pref., Japan - KC552038 KC552082 -
Tylopilus alpinus HKAS 55438 China - KF112404 - KF112538
Tylopilus ballouii Osmundson
1198
Thailand - EU430740 - EU434340
Fraser Island, Queensland,
Australia
- -
Tylopilus felleus AT2001011 Uppsala, Sweden - JQ327015
HKAS 90203 China - KT990545 - KT990913
MCVE98230 Italy JF908787 - - -
Tylopilus ferrugineus Erie Co., NY, USA - -
Tylopilus microsporus (T.
neofelleus)
Yunnan Prov., China - KF112450 KF112225 -
Tylopilus otsuensis HKAS 53401 Chenzhou, Hunan, China - KF112449 KF112224 -
Tylopilus plumbeoviolaceus NY, USA - KF030350 KF030439 -
Veloporphyrellus alpinus HKAS 57490 Yunnan Prov., China - KF112380 KF112209 KF112555
Xerocomellus chrysenteron Bavaria, Germany - KF030415 -
Xerocomellus zelleri REH 8724 Humbolt Co., CA, USA - KF030271
Xerocomus magniporus HKAS 59820 Yunnan, SW China - -
Xerocomus perplexus MB00-005 USA - JQ003702 KF030438 -
Xerocomus subtomentosus England, U.K. - KC215222 - -
Bavaria, Germany - JQ327035 KF030391
Zangia citrina China - - -
Zangia erythrocephala Nujiang, Yunnan, Prov., China - KF112414 -
Zangia roseola China - - JQ928595
Uncultured Boletaceae clone 47C_G1_
H9
Jonathan Dickinson State Park,
Hobe Sound, Florida, USA
- - -
clone 4C_G2_C3 Big Lagoon Start Park,
Pensacola, FL, USA
- - -
Table 2. Newly designed RPB1Bothia clade.
Primer name Sequence (5’ → 3’)
RPB1-32-F AGGCYGATATCGTGAGTCGC
RPB1-147-F CTCGAGYTATCGAGGCGT
RPB1-835-R ACCCTCRTCYTCRTCCTTGGG
RPB1-1091-R CCATCYACYGCTATACTCGG
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
347
0.97/98
1/100
1/90
- /0.87
1/95
1/100
1/99
1/77
1/84
0.9/87
0.97/79
0.94/ -
0.99/ -
0.97/75
0.95/ -
0.99/ -
0.98/ -
0.99/ -
0.96/73
0.99/79
0.97/85
0.98 /76
1/ -
0.96/73
0.97/ -
0.98/97
- /74
0.98 / -
0.96/ -
Phylloporus luxiensis HKAS 57036
Phylloporus luxiensis HKAS 75077
Phylloporus sp. HKAS 74680
Phylloporus sp. HKAS 74684
Phylloporus sp. HKAS 74679
Phylloporus imbricatus HKAS 54647
Phylloporus imbricatus HKAS 68642
Phylloporus yunnanensis HKAS 52225
Phylloporus brunneiceps HKAS 56903
Phylloporus rhodoxanthus MAN075
Phylloporus rhodoxanthus REH8714
Phylloporus rhodoxanthus JLM1808
Phylloporus rhodoxanthus MAN099
Phylloporus sp. HKAS 74681
Phylloporus sp. HKAS 74682
Phylloporus sp. HKAS 74683
Phylloporus pelletieri Pp1
Phylloporus pelletieri Q7199c
Phylloporus pelletieri K 128205
Phylloporus rubeolus HKAS 52573
Phylloporus sp. HKAS 74685
Phylloporus rhodoxanthus SAR 89.457
Phylloporus sp. HKAS 74687
Phylloporus leucomycelinus HKAS 74678
Phylloporus sp. HKAS 74688
Phylloporus rubrosquamosus HKAS 54542
Phylloporus rubrosquamosus HKAS 52552
Phylloporus maculatus HKAS 56683
Phylloporus bellus HKAS 56673
Phylloporus pachycystidiatus HKAS 54540
Phylloporus sp. HKAS 74689
Phylloporus gajari HKAS 76158
Phylloporus attenuatus HKAS 76168
Phylloporus rufescens HKAS 59722
Phylloporus catenulatus HKAS 76157
Hourangia cheoi HKAS 52269
Hourangia nigropunctata HKAS 76657
Phylloporus parvisporus HKAS 54768
Xerocomus subtomentosus Xs1
Xerocomus subtomentosus KM167686
Xerocomus perplexus MB00-005
Xerocomus magniporus HKAS 59820
Australopilus palumanus REH 6791
Royoungia boletoides Trappe 27456
Harrya chromapes HKAS 50527
Zangia erythrocephala HKAS 75046
Rossbeevera vittatispora MEL2128491
Rossbeevera griseovelutina holotype TNSF36989
Turmalinea persicina holotype KPM-NC 18001
Leccinellum aff. griseum KPM-NC 17831
Leccinellum crocipodium KPM-NC 18041
Octaviania nonae KPM-NC 1774
Octaviania decimae KPM-NC 17763
Octaviania tasmanica MEL2341996
Octaviania kobayasii KPM-NC 17783
Octaviania yaeyamaensis KPM-NC 17819
Leccinum versipelle KPM-NC 17833
Leccinum scabrum KPM-NC 17840
Spongiforma thailandica DED 7873
Borofutus dhakanus HKAS 73792
Retiboletus fuscus HKAS 59460
Retiboletus griseus Both sn
Sutorius eximius REH 8594
Sutorius aff. eximius HKAS 52672
Austroboletus gracilis 112/96
Austroboletus fusisporus HKAS 75207
Veloporphyrellus alpinus HKAS 57490
Fistulinella prunicolor REH 9502
Caloboletus inedulis MB 06044
Caloboletus firmus MB 06060
Suillellus amygdalinus 112605ba
Rubroboletus rhodosanguineus 4252
Neoboletus magnificus HKAS 74939
Boletus pulchriceps DS4514
Butyriboletus appendiculatus Bap1
Porphyrellus porphyrosporus KPM-NC 22667
Porphyrellus porphyrosporus KPM-NC 25017
Porphyrellus porphyrosporus MB 97023
Porphyrellus porphyrosporus HKAS 76771
Porphyrellus sp. HKAS 75078
Porphyrellus sp. HKAS 53366
Afrocastellanoa ivoryana Arora 126
Porphyrellus brunneus REH 9527
Strobilomyces strobilaceus Sf1
Imleria badia Xb2
Imleria badia SF119691
Imleria obscurebrunnea HKAS 52557
Boletus edulis Be3
Boletus variipes var. fagicola AH Smith 4249
Boletus semigastroideus PBM3076
Tylopilus felleus AT2001011
Tylopilus ferrugineus MB 06053
Tylopilus balloui REH 9467
Tylopilus microsporus HKAS 59661
Tylopilus otsuensis HKAS 53401
Tylopilus plumbeoviolaceus MB 06-056
Afroboletus luteolus 00-436
Xerocomellus zelleri REH 8724
Xerocomellus chrysenteron Xch1
Heliogaster columellifer KPM-NC 23012
Nigroboletus roseonigrescens GDGM 43238
Boletus rufomaculatus 4414
Baorangia pseudocalopus HKAS 75081
Lanmaoa carminipes MB 06-061
Lanmaoa asiatica HKAS 63592
Phylloporopsis boletinoides JBSD127413 Dominican Republic
Phylloporopsis boletinoides JBSD127411 Dominican Republic
Phylloporopsis boletinoides JBSD127412 Dominican Republic
Phylloporopsis boletinoides JBSD127414 Dominican Republic
Phylloporopsis boletinoides CORT 010991 Texas, USA
Phylloporopsis boletinoides F1118420 Florida, USA
Phylloporopsis boletinoides CORT 014483 Belize
Bothia castanella MB 03-053 MA USA
Bothia fujianensis HKAS 82694 Fujian CHINA
Solioccasus polychromus REH 9417 QLD AUSTRALIA
Phylloboletellus chloephorus XAL3388
Gymnogaster boletoides REH 9455
Pseudoboletus parasiticus Xps1
Buchwaldoboletus lignicola HKAS 76674
Chalciporus piperatus MB 04-001
Gyrodon lividus Gl1
0.05
Hourangia sp. HKAS 68178
Hourangia microcarpa HKAS 83763
1/100
1/98
1/100
1/100
1/100
1/100
1
/100
1/100
1/100
1/100
1/100
1/100
1
/100
1/100
1/98
1
/96
1/100
1/96
1/72
1/100
1/100
1/100
1/100
1/100
1/97
1/99
1/100
1/100
1/100
1/100
1/100
1/100
1/100
1/100
1/92
1/100
1/86
1/86
1/100
1/99
1/100
1
/100
1
/100
1
/92
Bothia clade
Fig. 1. Phylogeny of the Boletaceae based on a Bayesian and Maximum likelihood inference analysis of a combined matrix of two nuclear gene regions
(28S and TEF1-α
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348
ITS sequences were calculated using Geneious v. 11.1.4 (Kearse
et al.
TreeBASE (www.treebase.org) under ID S22978.
RESULTS
Molecular analysis
Both Bayesian and Maximum Likelihood analyses produced
comparable topologies and therefore only Bayesian trees with
BPP and MLB values are shown (Figs 1–3). The combined 28S/
TEF1-α and 28S/RPB1 dataset comprised 123 and 45 taxa,
In the 28S/TEF- α dataset (Fig. 1), all the P. bolenoides
is part of a larger clade that includes Bothia and Solioccasus
includes Bothia, Solioccasus and P. bolenoides as the Bothia
clade. Phylloporus clusters with Xerocomus and Hourangia
is distantly related to the Bothia clade. In the combined 28S/
RPB1 dataset (Fig. 2), P. bolenoides was strongly supported in
the Bothia Phylloporus clusters
with Xerocomus in a strongly supported clade (BPP = 1.0, MLB
hymenophore, Phylloboletellus, is resolved in an isolated and
Boletaceae (Figs 1, 2). In the ITS
analysis (Fig. 3), the newly generated sequences of P. bolenoides
sequence from Jonathan Dickinson State Park, Hobe Sound,
sequence from Big Lagoon State Park, Pensacola, FL, USA. Both
sequences were obtained from ectomycorrhizal samples on
Pinus clausa
of the ITS sequences of the Phylloporopsis
Fig. 2. Phylogeny of the Boletaceae based on a Bayesian and Maximum likelihood inference analysis of a combined matrix of two nuclear gene regions
(28S and RPB1
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Phylloporopsis gen. nov.
Editor-in-Chief
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E-mail:p.crous@westerdijkinstitute.nl
349
TAXONOMY
Phylloporopsis Angelini, A. Farid, Gelardi, M.E. Smith, Costanzo,
& Vizzini, gen. nov. MycoBank MB828149.
Etymology: the generic epithet refers to the morphological
Phylloporus.
Basidiomata
hymenophore, epigeal, evelate, medium-small sized;
hymenophore lamellate to subporoid with anastomosing and
interveined gills, strongly decurrent, beige to olive-cream or olive-
hymenophore and pileus context when injured or exposed; taste
smooth, ellipsoid-fusiform, spore wall cyanophilic; pleuro-, cheilo-
Phylloporus
analysis of the combined ITS, 28S, TEF1-α, and RPB1 sequences
the genus is unrelated to Phylloporus and close but separated
from Bothia and Solioccasus.
0.82/84
1/99
0.7/59
1/100
1/100
0.98/81
0.87/64
Bothia castanella NY8669 USA
Bothia castanella MB 03-067 USA
Bothia castanella NY28002 USA
Bothia castanella NY28003 USA
Bothia fujianensis HKAS 82693 China
Bothia fujianensis HKAS 82694 China
Solioccasus polychromus J. Trappe 15399 Australia
Tylopilus felleus MCVE98230 Italy
0.07
1/100
1/100
Phylloporopsis boletinoides Farid 617 Florida, USA
Phylloporopsis boletinoides FLAS-F-60407 Florida, USA
Phylloporopsis boletinoides FLAS-F-61158 Florida, USA
Phylloporopsis boletinoides FLAS-F-60413 Florida, USA
Phylloporopsis boletinoides F1118420 Florida, USA
uncultured Boletaceae clone 4C_G2_C3 Florida, USA
uncultured Boletaceae clone 47C_G1_H9 Florida, USA
Phylloporopsis boletinoides CORT014483 Belize
Phylloporopsis boletinoides JBSD127413 Dominican Republic
Phylloporopsis boletinoides JBSD127411 Dominican Republic
Phylloporopsis boletinoides JBSD127412 Dominican Republic
Phylloporopsis boletinoides JBSD127414 Dominican Republic
Bothia castanella MB 03-053 USA
Phylloporopsis
Bothia
Solioccasus
Fig. 3. Bayesian ITS phylogeny restricted to the clade including Phylloporopsis (Bothia
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Typus generis: Phylloporopsis bolenoides (A.H. Smith & Thiers)
Vizzini, Angelini, A. Farid, Gelardi, Costanzo, & M.E. Smith
Phylloporopsis bolenoides (A.H. Smith & Thiers) Vizzini,
Angelini, A. Farid, Gelardi, Costanzo & M.E. Smith comb. nov.
Basionym: Phylloporus bolenoides A.H. Smith & Thiers, Contr.
Monogr. North Amer. Species Suillus
Typus
Lake, east of Gainesville, solitary in deep sandy humus under
pines (Pinus spp.), low hammock, 31 Jul. 1958, H.D. Thiers
[MICH 11740 (holotype), SFSU 000741 (isotype
Basidiomata medium-small. Ontogenec development
gymnocarpic. Pileus
gradually fading with age and becoming beige-ochraceous to
LIII). Hymenophore
arcuate-decurrent, somewhat distant, undulate, shorter than
beige to pale cream-beige (Marguerite Yellow, Primrose Yellow,
due to mature spores, staining light blue (Pale Green-Blue Grey,
Spe (2.5–)2.7–4.5(–5.0)
× 0.4–0.7 cm, shorter than or as long as the pileus diameter at
or curved, cylindrical but slightly swollen towards the base, not
Plate II) in the upper third, concolourous with the pileus to
when pressed; rhizomorphs not observed. Context
air (Fig. 4H), especially above the tubes and eventually fading to
Odour Taste mild
Spore-print not
obtained. Macrochemical reacons
to dark brown on context and hymenophore, golden yellow on
4OH: staining purple-pink on pileus. FeSO4: slowly
pale yellowish-green to olive green on context, none elsewhere.
Basidiospores
3, inequilateral, very variable in
dimensions and versiform, cylindrical to fusiform or more
frequently ellipsoid-fusiform to broadly ellipsoid in side view,
ellipsoid to broadly ellipsoid in face view, smooth, apex rounded,
suprahilar depression although in some spores the depression
to very weakly dextrinoid, cyanophilic (Fig. 5E) and with a
Basidia (27–)33–54(–58)
basidioles subcylindrical, cylindrical-clavate to clavate, similar
in size to basidia. Cheilocysdia (42–)45–108(–118) × (8–)10–
ventricose-fusiform to sublageniform, less frequently cylindrical
to irregularly cylindrical, subclavate, sausage-like or peanut-like,
rarely mucronate or subcapitate, with rounded to subacute
Pleurocysdia
Pseudocysdia not recorded. Pileipellis
tending to be repent in the outermost layer and thus turning
yellow to yellowish-orange (inamyloid to weakly dextrinoid) in
pigments. Spipellis a layer of slender, parallel to subparallel
Fig. 4. Phylloporopsis bolenoides. A–F. Fresh basidiomata (A: JBSD127411, B: JBSD127412, C: JBSD127413, D: JBSD127414, E: CORT014483, F:
CORT010991). G. Detail of the lamellate hymenophore (JBSD127411). H. Detail of the context turning blue on exposure (JBSD127415). Bars = 1 cm.
Pictures: A–D, G–H by C. Angelini; E–F by T.J. Baroni.
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
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E-mail:p.crous@westerdijkinstitute.nl
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352
Fig. 5. Phylloporopsis bolenoides. A–B. Lignicolous basidiomata (A: JBSD127414, B: JBSD127414). C. Elements of the pileipellis (JBSD127412). D. Basidia
E.F. Basidiospores under SEM (JBSD127411). C–D
and longitudinally running, smooth walled, adpressed hyphae,
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353
fusiform, ventricose-fusiform, irregular cylindrical to subclavate
Lateral
spe stratum under the caulohymenium absent. Spe trama
composed of confusedly and densely arranged, subparallel
Hymenophoral trama Phylloporus-
mediostratum is concolorous with the lateral strata. Oleiferous
hyphae very common. Clamp connecons
Hyphal system
Ecology: usually under Pinus spp., occasionally in mixed Pinus
and Quercus forests, solitary to gregarious or less frequently
but also on logs or living trunks of pine trees.
Edibility: unknown.
Materials examined: Dominican Republic, La Vega Province, Jarabacoa,
specimens growing on soil under Pinus occidentalis, 21 Dec. 2013,
C. Angelini (JBSD127411, ANGE120 and MG709); ibid
specimens growing on soil under P. occidentalis, 22 Dec. 2013, C.
Angelini (JBSD127412, ANGE121 and MG710); ibid., a single mature
specimen growing on soil under P. occidentalis C.
Angelini (JBSD127413, ANGE551); ibid., several basidiomata in all
of P. occidentalis, 28 Nov. 2017, C. Angelini (JBSD127414, ANGE1007
and MG711); ibid., several basidiomata in all developmental stages
P. occidentalis, 23 Nov. 2017,
C. Angelini (JBSD127415, ANGE1013 and MG712). Belize, Cayo District,
Mountain Pine Ridge, Hidden Valley Inn property, near Lake Lolly Folly,
Jan. 2002, T.J. Baroni (9195 TJB = BZ745) (CORT014483). USA, Alabama,
Baldwin Co., Orange Beach, solitary in a sandy area with oaks nearby,
21 Jul. 1982, D.P. Lewis
Fig. 6. Phylloporopsis bolenoides. Drawings of the anatomical features (JBSD127412). A. Elements of the pileipellis. B.C.
Basidia. D.del.).
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354
hammock, 31 Jul. 1958, H.D. Thiers holotype);
Pinus sp., 11 Aug. 1985, N.S. Weber
line, 1.3 km E of C-731, 9 Sep. 2012, A.R. Franck 3125 (USF 273159);
Hillsborough Co., Violet Cury Nature Preserve, under Quercus sp. and
Pinus sp., 15 Jun. 2017, A. Farid
Pinus
N. Kraisitudomsookibid., under Quercus sp. and
Pinus M.E. Smith
M.E. Smith s.n. (USF 298023); Sarasota Co., Myakka Valley Ranches, on
soil near Pinus sp. and Quercus R.S. Williams
Pinus sp., 12
Jan. 1991, R. Singer (F3912) (F 1118420); Mississippi, Harrison Co.,
Pinus sp., 10 Sep. 1981, D.
Guravich
Cemetery, solitary in Pinus D.P. Lewis 3324
Pinus sp.
D.P. Lewis ibid., solitary on well-
decayed Pinus taeda stump, 13 Sep. 1979, D.P. Lewis 1982 (F 1089019);
T.J. Baroni (TJB 8172) (CORT010991).
Known distribuon: Eastern and south-eastern USA (Maine,
Delaware, New Jersey, Georgia, Florida, Mississippi, Alabama,
mycoportal.org/portal/index.php. Accessed on June 29, 2018),
in Central America reported from Belize and in the Greater
in tropical areas, western and southern limits yet to be
established.
DISCUSSION
Taxonomic circumscripon of P. bolenoides
features that characterize P. bolenoides.
reddish-orange then deep red or cinnamon-brown to cocoa brown
and becomes pale ochraceous-brown with age. The strongly
turns purplish-pink or reddish with ammonia. Anatomical traits
include the ellipsoid-fusiform, cyanophilic, smooth basidiospores,
Phylloporus-
Singer et al.et al.et al.
2007, Neves & Halling 2010, this study). The Dominican material
basidiospores, (11.1–)13.4 ± 1.27(–18.0) × (4.7–)5.8 ± 0.59(–8.2)
2.7 in Singer et al.
Santana et al.
& Halling 2010). This morphological variability likely represents a
species (Fig. 3).
was updated by Singer et al. (1990) to point out the inconsistent
samples found in Alachua County, Florida, near the type locality
bluing. A recent treatment on the boletes from eastern North
et al.
in P. bolenoides
et al. 1990).
Data obtained from top BLASTn results on GenBank and ITS
P. bolenoides with Pinus.
(clone 4C_G2_C3) were obtained from ectomycorrhizal root
samples from Pinus clausa. Florida material is typically found
under stands of Pinus spp., though occasionally in mixed Pinus
and Quercus forests. In the Dominican Republic P. bolenoides
was found with Hispaniolan pine (Pinus occidentalis) at high
Fagaceae occur in the Dominican Republic), despite several
trees (Figs 4D, 5A, B). The occurrence of the Belizean material
of P. bolenoides et al. 2007)
some neotropical sites. Previous studies have shown that
et al. 1985, Henkel et al.
2000). Examples of lignicolous growth of ECM Boletaceae have
been reported from the Americas, Europe, southeast Asia and
Australasia and include taxa such as those of the Boletellus
ananas complex and several Tylopilus species (Singer 1945a,
Corner 1972, Alessio 1985, Rayner et al. 1985, Henkel 1999,
et al. 1987, Henkel et al.
2000, 2012, Lindahl & Tunlid 2015).
Outside of the USA, P. bolenoides
restricted to Belize and the Dominican Republic but given its
throughout the neotropical mainland and the Caribbean where
pines occur.
Taxonomic and phylogenec relaonships of
Phylloporopsis to Bothia, Phylloporus, Phylloboletellus
and Phyllobolites
Phylloporopsis
Bothia
Bolenus castanellus) and the sequestrate Solioccasus
with S. polychromus)
Phylloporus sensu stricto (Figs 1–2). Based on the available data
there are no obvious shared morphological or ecological features
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
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355
the spore wall (Trappe et al.
TJB on B. castanella
synapomorphy known to date, though this feature has been only
Boletaceae
et al. (2013) established the
genus Solioccasus as a sister clade with Bothia, although they
did not address the shared morphological character between
the two genera. More extensive sampling is needed to elucidate
the taxonomic boundaries of the Bothia clade.
Though Phylloporopsis is Bothia,
Bothia
and radially stretched pores that are only slightly decurrent
et al. 2007). Conversely, in Phylloporopsis the basic structure
of the hymenophore is lamellate though anastomosing,
Bothia and blue in Phylloporopsis
and the spore print is pale brownish-yellow in Bothia while in
Phylloporopsis et al. 1990,
et al
Bothia is associated with Fagaceae (mostly Quercus
spp.) whereas Phylloporopsis forms ECM with pine trees (see
below). Finally, the geographic range of Bothia
overlapping with that of Phylloporopsis, since B. castanella (=
Bolenus squarrososides according to Halling et al. 2007) is
only found in eastern and south-eastern North America [Peck
B. squarrososides
Phylloporus squarrosoides
Xerocomus squarrosoides B.
squarrososidesB. squarrososides
et al.
Roody 2003, Watling 2008, Halling et al.
B. fujanensis is restricted
to south-eastern China (Fujian and probably Taiwan) (Chen et al.
1997, Zeng et al. 2015). These characters are a sound basis for a
Bothia castanella
P. bolenoides
B. castanella
(Halling et al.et al.
Phylloporopsis suggests that
the Boletaceae from poroid ancestors, viz. in Phylloporopsis,
Phylloporus and Phylloboletellus.
Phylloporopsis is morphologically similar to species of
Phylloporus
However, Phylloporus
golden-yellow lamellate hymenophore with the bright yellow
microscope (SEM) (although a number of tropical species have
et
al. 2010, 2012, Zeng et al.
trait that was considered unifying in Phylloporus is the pileus
surface that stains vivid blue to bluish-green with ammonia
Phylloporopsis. However, it
tested on species described from tropical Africa (Heinemann
1951), Malaysia (Corner 1970, 1974), China (Zeng et al. 2013, Ye
at al
a few taxa from tropical Central and South America, India and
Thailand (e.g. P. bulatus, P. manausensis, P. septocysdiatus, P.
castanopsidis
3 and NH4
2010, Neves et al. 2012, Pradeep et al. 2015). Accordingly, the
bluing of the pileus surface with ammonia cannot be considered
a synapomorphy within Phylloporus. Phylloporus is a large
assemblage encompassing nearly one hundred species so far as
yet been formally described), it appears to be widespread being
best represented throughout temperate and pantropical regions
of both hemispheres and is apparently most diverse in Australasia
Montoya & Bandala 1991, 2011, Halling et al. 1999, Watling & Li
et al. 2007, Neves & Halling 2010, Neves et
al. 2012, García-Jiménez 2013, Ye et al. 2014, Hosen & Li 2015,
2017, Pradeep et al. 2015). The neotropical P. centroamericanus
P. bolenoides but diverges,
apart from the generic discrepancies cited above, in the smaller
size (pileus 2–3 cm broad), pileus surface tending to become
with some eastern Asian species such as P. rubiginosus and P.
pachycysdiatus; the former is separated by the bright yellow
to orangish-yellow lamellae, context evenly yellowish and slowly
surface with ammonia, smaller spores [9.8–11.2(–13) × 3.5–4.9
the occurrence under Castanopsis and Dipterocarpus in Thailand
and south-western China (Yunnan Province) (Neves et al. 2012,
Ye et al.
size (pileus 3–5 cm in diameter), yellow hymenophore, cream-
Lithocarpus
southern and south-western China (Hainan and Yunnan Provinces)
(Zeng et al. 2013, Ye et al. 2014). There appears to exist a minor
P. bolenoides and the type species of the genus
Phylloporus, P. pelleeri, which is separated from the former by
the bright yellow to golden yellow hymenophore, bright yellow
ammonia on pileus surface, bacillate basidiospores under SEM
and the occurrence in temperate Europe (Pilát & Dermek 1974,
Breitenbach & Kränzlin 1991, Lannoy & Estadès 2001, Ladurner
& Simonini 2003, Watling & Hills 2005, Klofac 2007, Muñoz et al.
Boletaceae
is Phylloboletellus, a monotypic genus based on P. chloephorus.
This taxon was described from subtropical to tropical montane
Mexico (García-Jiménez et al. et al.
© 2018 Westerdijk Fungal Biodiversity Instute
Farid et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
1992, Bandala et al. 2004, García-Jiménez 2013). This species,
P. bolenoides by the subumbonate pileus,
yellowish basal mycelium, evenly yellow context discolouring
Boletus
short, broadly ellipsoid, longitudinally winged basidiospores
Petersen 1974, Pegler & Young 1981, Singer et al. 1992, Bandala
et al. 2004, Watling 2008). Moreover, the original material of
P. chloephorus was collected under Lauraceae, Myrtaceae,
Sapindaceae, Ulmaceae, etc., but the authors were unable to
the species was thought to be presumably saprotrophic (Singer
subsequently Bandala et al.
Tedersoo et al.
Phylloboletellus chloephorus var. mexicanus ad interim was
informally proposed by Singer et al. (1992) to circumscribe the
mexicanus with respect to the type
species were nearly inconsistent, thus it has never been validated
et al. 2011,
2014, Zeng et al.
P. chloephorus.
It is also worth nothing here the existence of the poorly
known genus Phyllobolites, which some authors have argued
may belong to the Boletaceae (Kirk et al. 2008, Magnago 2014).
Phyllobolites was erected based on Phyllobolites miniatus
from tropical northern South America (Guyana, Suriname,
et al. 1983, Henkel et al.
2012, Sulzbacher et al. 2013). This taxon has been originally
assigned to the broadly conceived family Paxillaceae, along with
unrelated agaricoid genera such as Omphalotus, Lampteromyces,
Hygrophoropsis, Neopaxillus and Ripartes
is easily delimited by the presence of a membranous ring deriving
Singer et al. 1983, Watling 2008). Since it has not been possible to
locate and re-examine the type specimen of P. miniatus from the
Rick was found by Singer to be referred to either Lennus (Singer
Tapinella or Pleurotus, the assignment of Phyllobolites to
the Boletaceae (Kirk et al. 2008, Magnago 2014) or Paxillaceae
et al. 1983, Neves &
Capelari 2007, Henkel et al. 2012) or even to the order Boletales
Phyllobolites
might belong in the order Gomphales near the genus Linderomyces
(a later synonym of Gloeocantharellus) (Watling, 2008). A modern
ACKNOWLEDGEMENTS
(MICH) for the loan of dried fungal materials. CA also wishes to thank
Ricardo G. García, Francisco Jiménez, Brígido Peguero, Yuley E. Piñeyro
and Alberto Veloz (Jardín Botánico Nacional Dr. Rafael M. Moscoso, Santo
Domingo, Dominican Republic) for their interest and encouragement in
technical help in taking SEM images. Mycological biodiversity research in
funding from the Faculty Research Program of the State University of New
York – College at Cortland for research work in the Gulf Coast region of
Arizona State University for his assistance with sequencing. Funding for
Ordway-Swisher Research grants (to MES).
REFERENCES
Alessio CL (1985). Boletus Dill. ex L. Fungi Europaei 2. Giovanna Biella,
Saronno, Italy.
Altschul SF, Gish W, Miller W, et al. (1990). Basic local alignment search
tool. Journal of Molecular Biology 215: 403–410.
Alvarado P, Moreno G, Vizzini A, et al. (2015). Atractosporocybe,
Leucocybe and Rhizocybe, three new clitocyboid genera in the
Tricholomatoid clade (Agaricales
species of Clitocybe and Lepista. Mycologia 107
Persoonia
18
North American boletes.
A color guide to the eshy pored mushrooms. Syracuse University
Press, Syracuse, USA.
Boletes of eastern North
America. Syracuse University Press, Syracuse, USA.
Boletales. Mycologia 98: 971–981.
Both EE (1993). The boletes of North America. A compendium
Breitenbach J, Kränzlin F (1991). Fungi of Switzerland. Vol. 3. Boletes
and agarics 1. Mykologia, Luzern, Switzerland.
Regensburger Mykologische Schrien 11
Chen C-M, Peng J-J, Yeh K-W (1997). The boletes of Taiwan (VIII).
Taiwania 42
Coker WC, Beers AH (1943). The Boletaceae of North Carolina. University
of North Carolina Press, Chapel Hill, USA.
Corner EJH (1970). Phylloporus Quél. and Paxillus Fr. in Malaya and
Borneo. Beihee zur Nova Hedwigia 20: 793–822.
Corner EJH (1972). Boletus in Malaysia. The Botanic Gardens,
Corner EJH (1974). Boletus and Phylloporus in Malaysia: further notes
Gardens’ Bullen Singapore 27
Darriba D, Taboada GL, Doallo R, et al.
Nature Methods
9: 772.
using the bootstrap. Evoluon 39: 783–791.
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
357
García-Jiménez J (2013). Diversidad de macromicetos en el estado
de Tamaulipas, México
Mexico.
et al.
boletáceos en México. Revista Mexicana de Micología 2
Basidiomycetes
and rusts. Molecular Ecology 2: 113–118.
Halling RE, Baroni TJ, Binder M (2007). A new genus of Boletaceae from
eastern North America. Mycologia 99
Halling RE, Mueller GM, Dallwitz MJ (1999). A new Phylloporus
(Basidiomycetes, Boletaceae) with a key to species in Colombia and
Costa Rica. Mycotaxon 63
Halling RE, Mueller GM (2005). Common Mushrooms of the Talamanca
Mountains, Costa Rica. Memoirs of the New York Botanical Garden
90. New York Botanical Garden Press, Bronx (NY), USA.
Heinemann P (1951). Champignons récoltés au Congo Belge par
Bullen du Jardin
Botanique de l’État à Bruxelles 21
Heinemann P, Rammeloo J (1987). Phylloporus (Boleneae). Flore
Illustrée des Champignons d´Afrique Centrale 13: 277–309.
Tylopilus from
Dicymbe forests of Guyana. Mycologia 91
Russulaceae from Guyana and Japan, with notes on their
ectomycorrhizal status, Mycologia 92: 1119–1132.
Henkel TW, Aime MC, Chin MML, et al. (2012). Ectomycorrhizal
fungal sporocarp diversity and discovery of new taxa in Dicymbe
monodominant forests of the Guiana Shield. Biodiversity and
Conservaon 26: 2195–2220.
Synopsis generum Agaricalium (Die Gaungstypen der
Agaricales)
Waber, Bern, Switzerland.
Hosen I, Li T-H (2015). Phylloporus gajari, a new species of the family
Boletaceae from Bangladesh. Mycoscience 56: 584–589.
Hosen I, Li T-H (2017). Two new species of Phylloporus from Bangladesh,
with morphological and molecular evidence. Mycologia 109: 277–
Jurgensen MF, Larsen MJ, Graham RT, et al.
in woody residue of northern Rocky Mountain conifer forests.
Canadian Journal of Forestry Research 17: 1283–1288.
Bolete Workshop – Studies in Boletology. Retrieved
Katoh K, Misawa K, Kuma K, et al. (2002). MAFFT: a novel method for
Nucleic Acids Research 30
Kearse M, Moir R, Wilson A, et al. (2012). Geneious Basic: an integrated
analysis of sequence data. Bioinformacs 28
Kirk PM, Cannon PF, Minter DW, et al. (eds.) (2008). Diconary of the
Fungi. 10th
europäischen Arten der Boletales mit röhrigem Hymenophor.
Österreichische Zeitschri für Pilzkunde 16: 187–279.
Knudsen H, Taylor AFS (2012). Xerocomus Quél. In: Funga Nordica –
Agaricoid, boletoid, clavarioid, cyphelloid and gastroid genera (2nd
ed.). Vol. 1 (Knudsen H, Vesterholt J, eds). Nordsvamp, Copenhagen:
228–233.
Ladurner H, Simonini G (2003). Xerocomus s.l. Fungi Europaei 8. Edizioni
Candusso, Alassio, Italy.
Lannoy G, Estadès A (2001). Flore Mycologique d’Europe 6 - Les
Bolets
Li YC, Feng B, Yang ZL (2011). Zangia, a new genus of Boletaceae
supported by molecular and morphological evidence. Fungal
Diversity 49: 125–143.
Li YC, Li F, Zeng NK, et al. (2014). A new genus Pseudoaustroboletus
(Boletaceae, Boletales) from Asia as inferred from molecular and
morphological data. Mycological Progress 13
New Phytologist 205:
1443–1447.
Magnago AC (2014). Taxonomia e sistemáca de Boletaceae (Boletales)
para o Brasil
Fungos, Algas e Plantas, Universidade Federal de Santa Catarina,
Proceedings
of the Gateway Compung Environments Workshop (GCE), 14
November 2010, New Orleans, LA: 1–8.
Montoya L, Bandala VM (1991). Studies on the genus Phylloporus in
new species and a new record. Mycotaxon 41: 471–482.
Montoya L, Bandala VM (2011). A new Phylloporus from two relict
Fagus grandifolia var. mexicana
forest. Mycotaxon 117: 9–18.
Xerocomus en la Peninsula Iberica.
Bolen de la Sociedad Micológica de Madrid 32: 249–277.
Murrill WA (1909). The Boletaceae of North America – I. Mycologia 1:
4–18.
Murrill WA (1914). American Boletes. Privately published, New York.
Neves MA (2007). Toward a revision of the genus Phylloporus (Boletaceae):
systemacs and phylogeny of species from various parts of the world,
Neves MA, Capelari M (2007). A preliminary checklist of Boletales from
Brazil and notes on Boletales
(SP) Herbarium, São Paulo, SP, Brazil. Sienbus Série Ciências
Biologicas 7
Neves MA, Halling RE (2010). Study on species of Phylloporus I:
Neotropics and North America. Mycologia 102: 923–943.
Neves MA, Henkel TW, Halling RE (2010). Phylloporus colligatus sp. nov.,
a new gilled bolete from Guyana. Mycotaxon 111: 143–148.
Neves MA, Binder M, Halling RE, et al. (2012). The phylogeny of selected
Phylloporus species, inferred from NUC-LSU and ITS sequences, and
Fungal Diversity
55: 109–123.
Nuhn ME, Binder M, Taylor AF, et al.
the Boleneae. Fungal Biology 117: 479–511.
Octaviania ivoryana (Boletaceae, Boletales), and the
proposal of a new genus, Afrocastellanoa. Mycologia 109: 323–332.
et al. (2007). Boletes from Belize
and the Dominican Republic. Fungal Diversity 27
Peck CH (1900). New species of fungi. Bullen of the Torrey Botanical
Club 27
Pegler DN, Young TWK (1981). A natural arrangement of the Boletales,
with reference to spore morphology. Transacons of the Brish
Mycological Society 76
Petersen RH (1974). Notes on cantharelloid fungi V. Some fungi
suspected of being gomphoid. The Journal of the Elisha Mitchell
Scienc Society 90: 53–54.
© 2018 Westerdijk Fungal Biodiversity Instute
Farid et al.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
358
Pilát A, Dermek A (1974). Hríbovité huby. Československé hríbovité a
sliziakovité huby (Boletaceae – Gomphidiaceae)
Pradeep CK, Vrinda KB, Varghese SP, et al. (2015). A new species of
Phylloporus (Agaricales, Boletaceae) from India. Phytotaxa 226:
overview. In: Developmental biology of higher fungi (Moore D,
Casselton LA, Wood DA, et al., eds). Cambridge University Press,
Cambridge: 1–40.
Rehner SA, Buckley E (2005). A Beauveria phylogeny inferred from
and links to Cordyceps teleomorphs. Mycologia 97: 84–98.
Rehner SA, Samuels GJ (1994). Taxonomy and phylogeny of Gliocladium
analysed from nuclear large subunit ribosomal DNA sequences.
Mycological Research 98
Brotéria, série
Botânica 5: 4–53.
Basidiomycetes eubasidii in Rio Grande do Sul - Brasilia.
5. Agaricaceae. Iheringia, série Botânica 8
Ridgway R (1912). Color standards and color nomenclature. Self-
published, Washington D.C.
Ronquist F, Teslenko M, van der Mark P, et al. (2012). MrBayes 3.2:
a large model space. Systemac Biology 61: 539–542.
Roody WC (2003). Mushrooms of West Virginia and the Central
Appalachians. University of Kentucky Press, Lexington.
Singer R (1938). Sur les genres Ixocomus, Bolenus, Phylloporus,
Gyrodon et Gomphidius. 2. Les Bolenus. Revue de Mycologie 3:
157–177.
Singer R (1942). Das System der Agaricales. II. Annales Mycologici 40:
1–132.
Singer R (1945a). The Boleneae of Florida with notes on extralimital
species. I. The Strobilomycetaceae. Farlowia 2: 97–141.
Singer R (1945b). The Boleneae of Florida with notes on extralimital
species. II. The Boletaceae (Gyroporoideae). Farlowia 2: 223–303.
Boleneae of Florida with notes on extralimital
species. IV. The lamellate families (Gomphidiaceae, Paxillaceae, and
Jugasporaceae). Farlowia 2
Singer R (1953). Type Studies on Basidiomycetes. VI. Lilloa 26: 57–159.
Monograph of South American Basidiomycetes, especially those
of the east slope of Andes and Brazil VI. The families Paxillaceae,
Gomphidiaceae, Boletaceae and Strobilomycetaceae. Nova
Hedwigia 7: 93–132.
Singer R (1970). Strobilomycetaceae (Basidiomycetes). Flora Neotropica
5. Hafner, New York.
Singer R (1978). Notes on bolete taxonomy – II. Persoonia 9: 421–438.
Singer R (1981). Notes on bolete taxonomy – III. Persoonia 11
The Agaricales in Modern Taxonomy. 4th ed. Koeltz
Basidiomycetes,
Agaricales) en relacion a las especies Mexacanas. Revista Mexicana
de Micología 4
Lilloa 22
ectomycorrhiza in Amazonian forests II. The Ectotrophically
Mycorrizal Fungi of the Neotropical Lowlands, Especially Central
Amazonia. Beihee Nova Hedwigia 77: 1–352.
Basidiomycetes of Costa Rica. III. The
genus Phylloporus (Boletaceae). Brenesia 22
Singer R, Ovrebo CL, Halling RE (1990). New species of Phylloporus
and Tricholomopsis from Colombia, with notes on Phylloporus
bolenoides. Mycologia 82: 452–459.
Boleneae of Mexico and
Central America IV. Beihee zur Nova Hedwigia 105
Singer R, Williams R (1992). Some boletes from Florida. Mycologia 84:
724–728.
A contribuon toward a monograph of
North American species of Suillus (Boletaceae). Privately published,
Ann Arbor.
Mycologia 28
discussions and a new subspecies. Mycologia 50
Snell WH, Dick EA (1970). The bole of northeastern North America. SH
Service Agency, J. Cramer, Lehre.
Bioinformacs 22
Sulzbacher MA, Grebenc T, Jacques RJS, et al. (2013). Ectomycorrhizal
fungi from southern Brazil – a literature-based review, their origin
Mycosphere 4
Xerocomus s.l. in the light of the present state of
knowledge. Czech Mycology 60
Tedersoo L, May TW, Smith ME (2010). Ectomycorrhizal lifestyle in
lineages. Mycorrhiza 20
Index Herbariorum: a global
directory of public herbaria and associated sta. New York botanical
Trappe JM, Castellano MA, Halling RE, et al. (2013). Australasian
sequestrate fungi 18: Solioccasus polychromus gen. & sp. nov., a
richly colored, tropical to subtropical, hypogeous fungus. Mycologia
105: 888–895.
Cryptococcus
species. Journal of Bacteriology 172
Watling R (2008). A manual and source book of the boletes and their
allies
Watling R, Hills AE (2005). Boletes and their allies - Boletaceae,
Strobilomycetaceae, Gyroporaceae, Paxillaceae, Coniophoraceae,
Gomphidiaceae Brish Fungus
Flora, Agarics and Bole. Vol. 1 (Henderson DM, Watling R eds).
HMSO, Edinburgh.
Watling R, Li T-H (1999). Australian boletes. A preliminary survey. Royal
Botanic Garden Edinburgh, Edinburgh.
Plant and Soil 90: 419–425.
White TJ, Bruns TD, Lee S, et al.
PCR protocols: a guide to methods and applicaons (Innis MA,
Gelfand DH, Sninsky J, et al., eds). Academic Press, San Diego: 315–
322.
et al.
the fungal family Boletaceae. Fungal Diversity 69: 93–115.
et al.
from China. Fungal Diversity 81: 25–188.
et al. (2014). The Genus Phylloporus
(Boletaceae, Boletales), from Mekong River Basin (Yunnan Province,
China). Chiang Mai Journal of Sciences 41: 1–13.
Zeng N-K, Cai Q, Yang ZL (2012). Corneroboletus, a new genus to
accommodate the southeastern Asian Boletus indecorus. Mycologia
104: 1420–1432.
© 2018 Westerdijk Fungal Biodiversity Instute
Phylloporopsis gen. nov.
Editor-in-Chief
Prof. dr P.W. Crous,Westerdijk Fungal BiodiversityInstitute, P.O. Box 85167, 3508 AD Utrecht, The Netherlands.
E-mail:p.crous@westerdijkinstitute.nl
359
Zeng N-K, Tang L-P, Li Y-C, et al. (2013). The genus Phylloporus
(Boletaceae, Boletales
DNA sequence analyses. Fungal Diversity 58: 73–101.
Zeng N-K, Su M-S, Liang Z-Q, et al. (2015). A geographical extension of
the North American genus Bothia (Boletaceae, Boletales) to East
Asia with a new species B. fujianensis from China. Mycological
Progress 14: 1015.
et al. (2015). Hourangia, a new genus of
Boletaceae to accommodate Xerocomus cheoi and its allied species.
Mycological Progress 14: 37.
