Investigations in the boletes (Boletaceae) of southeastern USA: four novel species and three novel combinations

Abstract

The Boletaceae is the largest family of fleshy fungi in the Boletales. Despite the extensive history of work in the Boletaceae in North America, novel species and genera are continually being described. Multigene molecular phylogenetic analyses of five loci were combined with thorough morphological studies to investigate the taxonomy of several boletes from the southeastern USA. Based on our results, we describe four new species: Aureoboletus pseudoauriporus, Cyanoboletus bessettei, Hemileccinum floridanum, and Xerocomellus bolinii. We also propose three combinations to reflect the results of our molecular analyses: Cyanoboletus cyaneitinctus comb. nov., a bolete that is widespread across the eastern USA, C. cyaneitinctus f. reticulatus, and Lanmaoa sublurida, a rarely-documented bolete that is so far known only from Florida.

Full text

Submitted 25 March 2021, Accepted 10 September 2021, Published XX October 2021
Corresponding Author: James R. Garey – e-mail – garey@usf.edu 1
Investigations in the boletes (Boletaceae) of southeastern USA: four
novel species and three novel combinations
Farid A1, Bessette AE2, Bessette AR2, Bolin JA3, Kudzma LV4, Franck AR5 and
Garey JR1*
1Herbarium, Department of Cell, Molecular, and Microbiology, University of South Florida, Tampa, Florida 33620
23109 William Penn Ct, Burlington, NC 27215
37340 Viale Sonata, Lake Worth, Florida 33467
437 Maple Ave., Annandale, NJ 08801
5University of Florida Herbarium, Florida Museum of Natural History, Gainesville, Florida 32611
Farid A, Bessette AE, Bessette AR, Bolin JA, Kudzma LV, Franck AR, Garey JR 2021 –
Investigations in the boletes (Boletaceae) of southeastern USA: four novel species, and three novel
combinations. Mycosphere X(X), X–X, Doi 10.5943/mycosphere/X/X/X
Abstract
The Boletaceae is the largest family of fleshy fungi in the Boletales. Despite the extensive
history of work in the Boletaceae in North America, novel species and genera are continually being
described. Multigene molecular phylogenetic analyses of five loci were combined with thorough
morphological studies to investigate the taxonomy of several boletes from the southeastern USA.
Based on our results, we describe four new species: Aureoboletus pseudoauriporus, Cyanoboletus
bessettei, Hemileccinum floridanum, and Xerocomellus bolinii. We also propose three combinations
to reflect the results of our molecular analyses: Cyanoboletus cyaneitinctus comb. nov., a bolete that
is widespread across the eastern USA, C. cyaneitinctus f. reticulatus, and Lanmaoa sublurida, a
rarely-documented bolete that is so far known only from Florida.
Keywords – Boletales – ectomycorrhizal – phylogeny
Introduction
The boletes of the southeastern USA are diverse but poorly studied. Perhaps the first taxonomic
work on the southeastern boletes began with Thomas Walter’s (Walter 1788) Boletus dimidiatus,
nom. illeg. Several more southeastern boletes were described by von Schweinitz (1822) and Berkeley
& Curtis (1853). Peck and Frost were both prolific with bolete studies in the late 1800s (Halling
1983, Both & Ortiz-Santana 2010). Murrill (1909) published the first monograph of the boletes of
North America, although after its publication, he described many more species of boletes (Halling
1986), especially from Florida (Weber 1961, Halling 1986). Coker & Beers (1943) published a
monograph of the boletes of North Carolina. Rolf Singer published a monograph on the boletes of
Florida (Singer 1945a, b, 1947), which treated species common to the southeastern USA, endemic to
Florida, and extralimital species from around the globe. Later, Murrill (1948) published a summation
of Florida boletes, one of the last broad treatments of the boletes of the region. Other works broadly
focused on boletes in the southeastern USA included Thiers (1963) and Grand (1970a, b, c). Both
(1993) published a compendium of all boletes described in North America, providing diagnostic
features as well as taxonomic notes on each species. Despite the extensive history and monographic
treatments, novel species of boletes from the southeastern USA are continually being described
Mycosphere X(X): X–X (2021) www.mycosphere.org ISSN 2077 7019
Article
Doi 10.5943/mycosphere/X/X/X

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(Singer & Williams 1992, Baroni 1998, Baroni et al. 1998, Ortiz-Santana et al. 2009, 2016, Frank et
al. 2017, Crous et al. 2019, Farid et al. 2020).
Molecular phylogenetic analyses have redefined our understanding of the boletes. Once
considered to consist of only a few genera, the Boletaceae has now increased to over 70 genera (Nuhn
et al. 2013, Wu et al. 2014, 2016). In part, this expansion is due to the recognition of sequestrate
(Yang et al. 2006, Smith et al. 2015, Castellano et al. 2016, Vadthanarat et al. 2018, Wu et al. 2018)
and new lamellate (Farid et al. 2018, Zhang & Li 2018) genera. This increase of genera is also due
to molecular phylogenetic analyses allowing taxonomists to better recognize synapomorphies, as
many of the traditional characters used to classify the boletes were homoplasic. The broad
relationships between genera are also better understood with analyses of molecular data. An analysis
of 290 operational taxonomic units (OTUs) across 59 genus-level clades by Wu et al. (2014) also
revealed six subfamily-level recognitions (Xerocomoideae, Leccinoideae, Boletoideae,
Austroboletoideae, Zangioideae, and Chalcioporoideae), although some genera did not resolve to any
of the known subfamilies (Solioccasus Trappe, Osmundson, Manfr. Binder, Castellano & Halling,
Bothia Halling, T.J. Baroni & Manfr. Binder, Gymnogaster J.W. Cribb, Baorangia G. Wu & Zhu L.
Yang, and Pseudoboletus Šutara), including one large grouping of genera (the Pulveroboletus group).
While much of the genus-level taxonomy has been explored in the Boletoideae, Xerocomoideae, and
the Pulveroboletus group (Murrill 1909, Halling et al. 2012, Vizzini 2014, Zhao et al. 2014, Gelardi
et al. 2015, Wu et al. 2016, Vadthanarat et al. 2019), many species-level taxa are still being described.
Boletes serve vital ecological roles as ectomycorrhizae of the primary forest trees (Quercus and
Pinus) of the southeastern USA, yet the extent of their diversity in this region is largely unknown.
The aim of this paper is to update our understanding of boletes in southeastern North America,
through multigene phylogenetic analyses. The name Boletus cyaneitinctus is resurrected for a species
closely related to Cyanoboletus pulverulentus (Opat.) Gelardi, Vizzini & Simonini. This paper
provides the first phylogenetic analyses of a rarely documented bolete, Suillellus subluridus Murrill,
which is transferred to Lanmaoa. We also describe four novel species, including one of
Xerocomellus, an uncommon species of Cyanoboletus, a species that resembles Hemileccinum
subglabripes (Peck) Halling, and one that resembles Aureoboletus auriporus (Peck) Pouzar. We also
generated protein-coding sequences from the epitype of Pulchroboletus rubricitrinus, as well as from
specimens of western Xerocomellus. Finally, we generated sequences from an herbarium specimen
of Exsudoporus floridanus from Florida and discuss the generic concepts of Exsudoporus and
Butyriboletus.
Materials & Methods
Sampling and morphological studies
Specimens were collected in situ between 2015–2020 and deposited at the University of South
Florida Herbarium (USF). Additional collections were obtained on loan from Florida Museum of
Natural History (FLAS) for study. Macroscopic descriptions were made using fresh basidiomes.
Micromorphological features were observed with a phase contrast microscope (AmScope, Irvine,
CA, USA). Distilled H2O, lactoglycerol, KOH, and Phloxine B were used to rehydrate and stain
sections (Singer 1986). Measurements were made at 1000 × with a calibrated ocular micrometer in
Piximètre 5.9 R 1532 (http://piximetre.fr). Basidiospore dimensions are reported as length by width,
with each measurement reported as the minimum, the average minus the standard deviation, the
average plus the standard deviation, and the maximum. Spore dimensions are followed by the number
of spores counted, N, and the average quotient mean, Q, where Q is the average length divided by
the average width. Scanning Electron Microscopy (SEM) was performed at the Electron Microscopy
Core Facility at the University of South Florida on an Aquila Hybrid Scanning Electron Microscope
(Topcon, Tokyo, Japan).
DNA Extraction, PCR amplification, and sequencing
Genomic DNA was isolated as described in Farid et al. (2017). A subset of the samples was

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extracted using the NucleoSpin Plant II Kit (Macherey-Nagel Inc. Bethlehem, Pennsylvania, USA).
Portions of five gene regions were targeted for phylogenetic analysis: nuc rDNA internal transcribed
spacer ITS1-5.8S-ITS (ITS), nuc 28S rDNA (28S), RNA polymerase II subunit 1 (RPB1), RNA
polymerase II subunit 2 (RPB2), and translation elongation factor 1-alpha (TEF1) were amplified
according to Farid et al. (2019). The primer pair ITS1-F/ITS4 (White et al. 1990). Gardes & Bruns
1993) were used to amplify ITS, LR0R/LR7 (Vilgalys & Hester 1990) for 28S. The bolete-specific
primer pairs EF1-BF1/EF1-B-R, RPB1-B-F/RPB1-B-R, and RPB2-B-F1/RPB2-B-R (Wu et al.
2014) were used to amplify TEF1, RPB1, and RPB2, respectively. Crude PCR product was purified
and sequenced at the DNA laboratory at Arizona State University with a 3730 DNA Analyzer
(applied Biosystems, Carlsbad, CA, USA) using the same PCR primers for amplification, and
additionally the internal 28S primers LR5 and LR3R were used (Vilgalys & Hester 1990).
A subset of samples (JAB 95 and JAB 80) was obtained using a nested PCR method. First, the
primer pair gRPB1-Af/fRPB1-Cr (Matheny et al. 2002) were used to amplify a portion of the RPB1
gene; PCR products were then diluted in nanoPure H2O in a 1:100 ratio used in a second hemi-nested
PCR using one of the original primers gRPB1-Af or fRPB1-Cr paired with an internal primer chosen
from either RPB1-B-F or RPB1-B-R or one of two novel Boletales specific primers (Table 1).
Table 1 Primer design Boletales-specific RPB1 primers
Primer name
Sequence (5’ → 3’)
RPB1mexF1bol
CGRCATGTYCGCGATCC
RPB1mexR2bol
GGWTCRTCAGYTTCGCA
Alignments, model selection, and phylogenetic analyses
A multi-locus phylogeny consisting of ITS, 28S, RPB1, RPB2, and TEF1. Alignments of each
locus were made in R (R Core Team 2017) using MAFFT v. 7.471; alignments of rDNA used the
predicted secondary structure to improve the alignment. Gblocks v. 0.91b (Katoh & Standley 2013)
was used to remove ambiguous regions of the resultant alignments to improve phylogenetic
inference. Models were selected for each locus using jModelTest 2.1.10 (Guindon & Gascuel 2003,
Darriba et al. 2012). Bayesian information criterion models were selected for each partition, though
we report all the models selected (Table 2). The resultant alignments were combined in Sequence
Matrix (http://www.ggvaidya.com/taxondna/), with taxa missing target loci encoded as missing data
(Felsenstein 2004). Seventeen genera from the Boletaceae were included in the phylogenetic analyses
(Fig. 1): Aureoboletus Pouzar, Hemileccinum Šutara, Pulchroboletus Gelardi, Vizzini & Simonini,
Heimioporus E. Horak, Alessioporus Gelardi, Vizzini & Simonini, Xerocomellus Šutara,
Nigroboletus Gelardi, Vizzini, E. Horak, T.H. Li & Ming Zhang, Hortiboletus Simonini, Vizzini &
Gelardi, Boletus L., Baorangia G. Wu & Zhu L. Yang, Cyanoboletus Gelardi, Vizzini & Simonini,
Lanmaoa G. Wu & Zhu L. Yang, Butyriboletus D. Arora & J.L. Frank, Suillellus Murrill,
Gymnogaster J.W. Cribb, Chalciporus Bataille, and Buchwaldoboletus Pilát.
Phylogenetic analyses were conducted using the CIPRES Gateway server V3.3 (Miller et al.
2010). Maximum likelihood (ML) was conducted with RAXML-HPC 8.2.10 (Stamatakis 2014)
using 1000 non-parametric bootstrap replicates (BS) and a partitioned model. Bayesian inference
(BI) was conducted with MrBayes 3.2.6 on XSEDE platform of the CIPRES Science Gateway server
(Ronquist et al. 2012). Four Markov chain Monte Carlo simulations were run for ten million
generations, sampling trees every thousand generations. Chain convergence was determined using
Tracer V1.6 (Rambaut et al. 2018). The first 25% were discarded as burn-in, and a majority rule
consensus tree was computed to obtain estimates for Bayesian posterior probabilities (BPP). BI trees
were visualized in Figtree (Rambaut 2007) and exported into Inkscape, where bootstrap values were
added to node labels. BPP above 0.90 and bootstrap values above 70% were reported. Alignment and
phylogenetic trees were uploaded to http://www.treebase.org/ (submission ID 27951).

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Table 2 Models selected for each locus using different model strategies in jModelTest 2.1.10.
Abbreviations: AICc = Akaike information criterion. BIC = Bayesian information criterion. DT =
Decision theory. GTR = Generalized time reversible model. HKY = Hasegawa, Kishino and Yano
1985 model. K80 = Kimura' s two parameter model. SYM = Symmetrical model. I = Invariant.
G = Gamma
Model Strategy
Locus
ITS
28S
RPB1
RPB2
TEF
AICc
GTR+I+G
GTR+I+G
HKY+I+G
SYM+I+G
HKY+I+G
BIC
GTR+I+G
GTR+I+G
K80+I+G
K80+I+G
HKY+I+G
DT
HKY+I+G
GTR+I+G
K80+I+G
K80+I+G
HKY+I+G
Figure 1 – Phylogram generated from MrBayes based on ITS, 28S, RPB1, RPB2, and TEF1 sequence
data. Nodes labeled with PP (≥ 0.90) followed by bootstrap replicate support (≥ 70). Colors represent

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distinct genera. Specimens with molecular data generated in this study are bolded. Inset phylogeny
depicts portion of phylogeny shown in figure.
Figure 1 – Continued.

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Figure 1 – Continued.

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Figure 1 – Continued.

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Results
Phylogenetic analyses
The final dataset consisted of 305 specimens comprising 141 ITS, 234 28S, 140 RPB1, 165
RPB2, 216 TEF1 sequences (Supplementary Table 1). A total of 143 sequences were generated for
this study. The six species from this study were distributed across five genera. One species of
Aureoboletus forms a strongly supported clade (0.96 BPP, 96 bootstrap replicate support), with
somewhat strong support (0.91 BPP, 94 bootstrap replicate support) as a sister clade with
Aureoboletus auriporus (Peck) Pouzar. A strongly supported clade in Hemileccinum with somewhat
strong support (0.96 BPP, but <70 bootstrap replicate support) was sister to a clade of Hemileccinum
subglabripes (Peck) Halling. In Xerocomellus, a strongly supported clade was sister to an unnamed
Xerocomellus sp. (HKAS 56311) from China. Three specimens of Nigroboletus roseonigrescens
Gelardi, Vizzini, E. Horak, T.H. Li & Ming were strongly supported as basal to all Xerocomellus
sequences included in the analyses. Two species in Cyanoboletus were recovered in the analyses.
This first Cyanoboletus species is in a strongly supported clade (1 BPP, 98 bootstrap replicate
support) containing Cyanoboletus pulverulentus s.str., and Cyanoboletus sinopulverulentus (Gelardi
& Vizzini) Gelardi, Vizzini & Simonini, although C. sinopulverulentus did not receive strong support
as sister to either of these species. The second Cyanoboletus species formed a strongly supported
sister clade to an unnamed Cyanoboletus sp. (HKAS 76850) from China, and a clade containing
Cyanoboletus instabilis (W.F. Chiu) G. Wu & Zhu L. Yang. A species of Lanmaoa formed a strongly
supported sister clade to Lanmaoa roseocrispans A.E. Bessette, A.R. Bessette, Nuhn & Halling.
Pulchroboletus rubricitrinus (Murrill) Farid & A.R. Franck, which was strongly supported as a sister
clade to Pulchroboletus roseoalbidus (Alessio & Littini) Gelardi, Vizzini & Simonini, was consistent
with the results from the nucDNA analysis in Farid et al. (2017). Our collection of Exsudoporus
floridanus formed a strongly supported clade with Exsudoporus floridanus from Belize (1.0 BPP,
100 bootstrap replicate support), while the Exsudoporus clade was strongly supported as sister to
Butyriboletus (1.0 BPP, 0.96 bootstrap replicate support).
Aureoboletus pseudoauriporus J.A. Bolin, A.R. Bessette, A.E. Bessette, L.V. Kudzma, A. Farid &
J.L. Frank sp. nov. Figs 2, 10D–F
MycoBank number: MB840856; Facesoffungi number: FoF 10467
Etymology – The epithet pseudoauriporus is from the Latin “pseudo” = false in reference to
this bolete so closely resembling, but differing from, Aureoboletus auriporus.
Typification – USA, Florida, Palm Beach County, Jupiter, Abacoa Natural Area, 1 Mar 2019,
J.A. Bolin 320 (holotype USF 301510).
Diagnosis – Medium-sized basidiocarps with a glabrous, non-viscid pinkish tan unchanging
pileus that becomes tan with age, or sometimes retains pinkish tones. The hymenophore is bright
yellow when young, becomes darker yellow and then dingy yellow with age, and does not stain when
bruised or cut. The stipe is typically longitudinally striate for one-third or more of its length.
Basidiospores measure (14-)15–17(-18) × 5-6.5 µm.
Description – Pileus 5–8.5 cm broad, convex at first, remaining so well into maturity; surface
glabrous, color variable, pinkish to pinkish red or pinkish tan, usually losing pinkish tones when
mature, unchanging when bruised, tastes acidic; margin incurved, even or narrowly sterile; staining
pale yellow-orange then fading to light brown with the application of KOH, pale blue-green fading
quickly or slowly with NH4OH, slowly staining light greenish gray or negative with FeSO4. Context
white, unchanging or faintly and slowly turning pink or light yellow near the hymenium; staining
yellow-orange with KOH, slowly light greenish gray or negative with NH4OH, and light blue-green
or negative with FeSO4; odor and taste not distinctive. Hymenophore tubulose, bright yellow when
young, becoming darker yellow and then dingy yellow with age, not staining when bruised or cut;
pores rounded, 1–2 per mm; tubes 4–12 mm deep. Stipe 4–6 cm long, 8–12 mm at the apex, 1–1.4
cm thick at the base, typically equal or slightly enlarged downward, sometimes with a pinched base;
surface typically dry but viscid when wet, typically longitudinally striate for one-third or more of its

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length, whitish, sometimes with pale pink tones, not staining when bruised; context white, firm and
woody toward the base, often staining faintly pinkish; with white basal mycelium.
Basidiospores light to medium brown in fresh deposit, (14–)15–17(–18) × 5–6.5 µm, n = 30,
Q = 2.79, elliptical in face view, inequilateral in profile, thick-walled, smooth, lacking an apical pore,
yellow-brown in KOH or Melzer’s. Basidia 25–38 × 8–13 µm, clavate, 2-sterigmate, hyaline in KOH
or Melzer’s. Basidioles 12–23 × 6.5–8 µm, clavate, thin-walled, hyaline in KOH or Melzer’s.
Hymenial cystidia 30–50 × 10–15 µm, cylindrical, sometimes with a capitate to capitulate apex.
Hymenophoral trama boletoid, with lateral elements, 4–12 µm wide, moderately divergent, hyaline
in KOH or Melzer’s. Pileipellis an ixotrichoderm, terminal elements 7–22 µm wide, highly variable,
thin-walled, smooth, hyaline in KOH, with golden yellow contents in Melzer’s. Pileus trama hyphae
loosely interwoven, highly variable, 6–32 µm wide, smooth, thin-walled, hyaline in KOH or
Melzer’s. Stipitipellis mostly parallel, slightly interwoven, 4–12 µm wide, hyaline in KOH or
Melzer’s, with fascicles of clavate or fusiform caulocystidia. Caulocystidia of two types; clavate, 24–
42 × 12–22 µm, with yellowish contents in KOH, thin-walled, smooth; fusiform 32–39 × 8–12 µm,
hyaline in KOH, thin-walled, smooth. Stipe trama interwoven, 6–13 µm wide, hyaline in KOH or
Melzer’s, thin-walled, smooth. Clamp connections absent.
Figure 2 – Field photograph of Aureoboletus pseudoauriporus. A J.A. Bolin 488. B J.A. Bolin 157.
C J.A. Bolin 124. D J.A. Bolin 130. Photo credit: J.A. Bolin.

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Habit, Habitat & Distribution – solitary or scattered in sandy soil with oak in a scrubby flatwood
community; known from central Florida, distribution limits yet to be determined.
Material examined – USA, Florida, Hillsborough County, Brandon, S of Camden Visconti
entrance pond, adjacent to canal, 27°55'27.9"N 82°20'22.9"W, 5 Oct 2016, A. Farid 501 (USF
288287); Tampa, Violet Cury Nature Preserve, 4 Jun 2017, A. Farid 592 (USF 301502); Tampa,
Trout Creek Nature Preserve, Xeric hammock beneath Quercus geminata, 3 May 2019, A. Farid 919
(USF 301507); Lake County, Lake Louisa State Park, Clermont, 24 Oct 2019, J.A. Bolin 448 (USF
301492); Miami-Dade County, Everglades National Park, 5 Jul 2019, A. Farid 959 with A.R. Franck
and R.E. O’Donovan (EVER 144770); Palm Beach County, Frenchman’s Forest natural Area, 28
May 2018, J.A. Bolin 167 (USF 301497); Hypoluxo Scrub Natural Area, Lantana, 21 Nov 2017, J.A.
Bolin 80 (USF 301487); ibid., 70 Sep 2019, 7 Sep 2019, J.A. Bolin 106 (USF 301489); ibid., 6 Nov
2017, J.A. Bolin 130, (USF 301493); Jupiter, Abacoa Natural Area, 1 Mar 2019, J.A. Bolin 320
(holotype USF 301510); ibid., 13 Aug 2019, J.A. Bolin 418 (USF 301483).
Notes – This species is a part of a cryptic species complex. It greatly resembles Aureoboletus
auriporus (Peck) Pouzar, and its distribution limits are yet to be established. Aureoboletus auriporus
differs from A. pseudoauriporus by the lack of longitudinal striations on the stipe. The pileus of A.
auriporus is reported to turn red with the application of NH4OH (Baroni 2017). The spore size of A.
auriporus was not originally reported in the protologue, though Peck (1889) later provided an
expanded description and reported the spores as 7.5–10 × 4–5 µm. Both (1998) studied the type
specimen, obtaining a spore size of 9.8–15.5 × 3.96–5.75 µm, with a mean dimension of 13.15 × 4.73
µm, Q = 2.12–3.39, Qm = 2.75. Both (1998) also provided a description based on collections
primarily from New York and Rhode Island, but also included a specimen from Tennessee, and did
not include the type specimen. The spores reported were slightly larger than the type, at 11.0–16.05
× 4.4–6.38 µm, mean dimension 14.36 × 5.19 µm, and the spore quotient was similar, at Q = 2.2 –
3.19, Qm = 2.78. The spores of A. pseudoauriporus are somewhat larger, at (14–)15–17(–18) × 5–
6.5 µm, × = 16.45 × 5.92 µm, and the spore quotient is nearly identical, at Q = 2.79.
Aureoboletus viridiflavus Coker & Beers ex Klofac is a similar species, and has been treated
as a synonym of A. auriporus in the past (Singer 1947, Both 1998), which differs primarily by the
pileus colors, which was described as “olivaceous gold with reddish areas”, the pileus when young
tomentose-felted, less viscid, a lack of distinctly projecting margin, the hymenophore longer, to 17.5
mm (4–12 mm in A. pseudoauriporus), and the stipe, which bruises “brick red” and is not viscid
(white, sometimes with pale pink tones, and not bruising in A. pseudoauriporus). The spore size is
similar to A. auriporus, reported as 11.5–15(–16.6) × 4–5 µm in the protologue. Aureoboletus
pseudoauriporus has somewhat longer and wider spores, measuring (14–)15–17(–18) × 5–6.5 µm.
Aureoboletus subacidus (Murrill ex Singer) Pouzar is a somewhat similar species that shares reddish
tones in the pileus, citrine yellow tubes, a whitish stipe, occurs in Florida, and is associated with
Quercus spp. It can readily be distinguished from A. pseudoauriporus by the presence of the floccose
yellow velar remnants left on the upper portion of the stipe, the scrobiculate pileus, and the yellow
pileal context (Singer 1947).
So far, A. pseudoauriporus is the only species in the complex known from Florida. Although
A. pseudoauriporus has been observed in southeastern Georgia (USA) by the authors, no collections
were made. Aureoboletus innixus (Frost) Halling, A. R. Bessette & A. E. Bessette is similar but it
has a dry, somewhat velutinous, dull reddish-brown pileus, and lacks longitudinal striations on its
stipe. Aureoboletus roxanae (Frost) Klofac has whitish pores when young which eventually become
pale yellow, and a yellow to pale orange-yellow stipe with a distinct dull orange zone at the apex.
Cyanoboletus bessettei A.R. Bessette, L.V. Kudzma, & A. Farid sp. nov. Figs 3, 10G–I
MycoBank number: MB 840857; Facesoffungi number: FoF 10466
Etymology – The epithet bessettei honors American mycologist, Alan E. Bessette.
Typification – USA, South Carolina, Berkeley County, Francis Marion National Forest, State
Route 402, approximately 1.25 mi. north of Huger, under oak and pine, 17 Sep 2016, A.R. Bessette
ARB1393 (Holotype USF 301500).

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Diagnosis – Medium-sized basidiocarps with a dry, reddish brown to buffy brown pileus and a
reddish-brown stipe with a pale-yellow apex and white basal mycelium. The hymenophore surface
is pale yellow and stains blue-green then olive when bruised. It has pale yellow context that stains
blue-green then slowly turns peach to dull pinkish orange when exposed. The basidiospores measure
(8–)9–11(–12) × 3.5–5 µm and are narrowly ovate to subelliptic. It fruits on the ground with oak and
pine during fall.
Description – Pileus 2.7–8 cm broad, convex with an incurved margin that remains into
maturity; surface subtomentose to nearly glabrous, dry, buffy brown overall when very young,
becoming paler toward the margin and retaining darker brownish coloration on the disc at maturity,
staining blue-green then dark olive-green and finally brown when bruised; margin with a narrow
band of sterile tissue, sometimes undulating or lobed in age; context pale yellow, staining blue-green
then slowly turning peach to dull pinkish orange when exposed; odor unpleasant, odd, chemical-like;
taste slightly acidic or not distinctive. Cuticle stains dark amber with the application of KOH, pale
olive with FeSO4, and amber with an expanding blue-green outer ring with NH4OH. Context stains
yellow, then pale orange with the application of KOH or NH4OH and is negative with FeSO4.
Hymenophore tubulose, pale yellow, staining blue-green, then olive when bruised; pores angular to
irregular, 2–3 per mm; tubes 4–8 mm deep. Stipe 2.5–4 cm long, 1–2 cm thick, nearly equal or flaring
at the apex, pinched at the base; surface longitudinally striate, dry, distinctly pale yellow at the apex,
reddish brown below, with white basal mycelium, staining blue-green then reddish-brown; context
pale yellow, slowly staining blue-green at the apex, then becoming bright chrome yellow.
Figure 3 – Field photograph of Cyanoboletus bessettei (ARB 1393). Photo credit: A.R. Bessette.
Basidiospores olive-brown in fresh deposit, (8–)9–11(–12) × 3.5–5 µm, n = 30, Q = 2.30,
narrowly ovate to subelliptic in face view, obscurely inequilateral in profile, thin-walled, smooth,
lacking an apical pore, yellowish in KOH or Melzer’s, inamyloid; spores sometimes collapsing when
mounted in Melzer’s. Basidia 23–36 × 5.5–9 µm, mostly clavate, few cylindro-clavate, (2)4-
sterigmate, hyaline in KOH, grayish yellow in Melzer’s. Basidioles 19–31 × 5–8.5 µm, clavate.

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Hymenial cystidia abundant, sometimes in fascicles, 36–51 × 8–11 µm, ventricose-rostrate, some
with an elongated neck, thin-walled, smooth, yellowish in KOH, non-reactive in Melzer’s.
Hymenophoral trama boletoid, with lateral elements, 3.5–7 µm, wide, moderately divergent, hyaline
to grayish yellow in KOH or Melzer’s. Pileipellis a tangled layer of repent tubular hyphae, 3.5–5.8
µm wide, with cylindrical, rounded end cells, thin-walled, smooth, hyaline to grayish yellow in KOH,
yellowish in Melzer’s. Pileus trama hyphae loosely interwoven, 4.5–13 µm, smooth, thin-walled,
hyaline in KOH, yellowish in Melzer’s. Stipitipellis hyphae mostly parallel, slightly interwoven, 3–
6.5 µm, hyaline in KOH, yellowish in Melzer’s. Stipe trama subparallel, interwoven, 5.2–14 µm,
hyaline in KOH, yellowish in Melzer’s. Caulocystidia none observed. Clamp connections absent.
Habitat and Distribution – Scattered or in groups in sparsely grassy areas and sandy soil in
association with Quercus and Pinus in fall (September), in southeastern United States, known only
from South Carolina.
Notes – The combination of distinctive staining reactions and very small spore size is distinct.
To date, it is known only from the type location in southeastern South Carolina, United States.
Caloboletus inedulis (Murrill) Vizzini is superficially similar with a pale yellow hymenophore that
becomes olive yellow at maturity and stains dark blue then brownish when bruised. It has a reticulated
pale-yellow stipe that may have pinkish tones at the apex and/or pinkish tints below, and bitter tasting
context.
Cyanoboletus cyaneitinctus (Murrill) A. Farid, A.R. Franck & J.A. Bolin comb. nov.
Figs 4, 5A–G, 10A–C
MycoBank number: MB 840858; Facesoffungi number: FoF 10465
Basionym – Ceriomyces cyaneitinctus Murrill, Lloydia 6: 225 (1943).
Synonyms – Boletus cyaneitinctus (Murrill) Murrill, Lloydia 6: 228 (1943).
Typification – USA, Florida, Alachua County, Gainesville, Kelley’s Hammock, 3 Aug 1938,
West and Murrill s.n. (holotype FLAS-F-17986); Hillsborough County, Tampa, Learning Gate
Community grounds, 4 May 2019, A. Farid 920 (epitype here designated USF 301499).
= Boletus mutabilis Morgan, J. Cincinnati Soc. Nat. Hist. 7: 6 (1884), nom. illegit., Art. 53.1.
Diagnosis – Brownish or rarely reddish pulvinate pileus, bright yellow hymenophore, stipe,
and context, all surfaces rapidly and brilliantly cyanescent.
Description – Pileus 3–8 cm wide, pulvinate or convex when young becoming broadly convex
at maturity, bister, umber, mahogany, and dark brown overall, rarely entirely red in the pileus,
glabrous to tomentose, tacky when wet, sometimes rimulose at maturity, blackening instantly where
handled. Hymenophore tubulose, yellow, darkening to a gold color when mature, tubes 5–20 mm
long, bluing instantly and strongly when handled; pore mouths subangular when mature, 0.5–1 mm
in diameter. Stipe 3–6 × 0.5–2 cm, equal to ventricose, bright yellow, smooth or sometimes reticulate
on the upper third, sometimes with flushes of reddish to brownish-red floccons, particularly towards
the base of the stipe, bluing instantly and strongly when handled, basal mycelium white to yellowish
white. Context concolorous with stipe surface, often with red pigments at the very base of the stipital
context, blueing instantly and strongly, fading to pale yellow. KOH on pileus dark maroon to black,
red elsewhere; FeSO4 negative, erasing blue stains from flesh.
Basidiospores (11)11.5–15(16) × 4–6 μm, n = 30, Q = 2.4, fusiform, sometimes with a
suprahilar depression present. Basidia 25–50 × 8–10 µm, 4-spored, thin-walled, hyaline, clavate to
pyriform; sterigmata 1–2 μm, occasionally pigmented like pleruocystidia. Basidioles similarly sized
and shaped. Pleurocystidia 30–60 × 7–10 µm, fusoid to ampullaceous, hyaline or sometimes
encrusted. Cheilocystidia similar to pleurocystidia. Pilepellis a trichodermium of strongly
interwoven, filamentous, sinuous, rarely branched hyphae, erect or repent in most of the terminal
elements, collapsing into a cutis, terminal elements cylindrical, apices rounded or somewhat pointed,
20–70 × 5–10 μm, smooth-walled, inamyloid, hyaline to golden-yellow or somewhat brownish in
water and 5% KOH. Clamp connections absent.
Habitat and Distribution – Basidiomes typically occurring singly or more rarely gregariously,
widely distributed in eastern North America.

13
Material examined – USA, Florida, Alachua Co., Gainesville, 2 Oct 1949, W.A. Murrill s.n.
(FLAS F16163); ibid., lawn under pecan [Carya illinoinensis], 7 Nov 1947, W.A. Murrill s.n. (FLAS
F40835); ibid., Kelley’s Hammock, 3 Aug 1938, West and W.A. Murrill s.n. (holotype FLAS
F17986); ibid., yard at 936 NW 30th Ave., 9 Aug 1980, G.L. Benny s.n. (FLAS F52704); ibid., lawn
under laurel oak [Quercus laurifolia], 1 Aug 1947, Murrill s.n. (FLAS F19093); ibid., shaded yard,
6 Nov. 1950, R. Bennett s.n. (FLAS F59706); ibid., lawn der hardwoods, 13 Oct 1950, R. Bennett
s.n. (FLAS F19647); ibid., 19 × 1950, R. Bennett s.n. (FLAS F 40863); ibid., lawn on 18th block of
NW 11 place, Sept 12 1968, J. Kimbrough s.n. (FLAS F48020); ibid., under large live oak [Quercus
virginiana] 10 mi. SE of Gainesville, on Palatka Rd., 2 Nov 1947, G.F. Weber s.n. (FLAS F40837);
Hillsborough Co., Alafia River State Park, 17 Jul 2018, J. Bolin 177 (USF 300090); Hillsborough
County, Tampa, University of South Florida Tampa Campus, entrance area off of Leroy Collins
Boulevard, 11 Jun 2016, A. Farid 340 (USF 288424); USF campus, 22 May 2018, Franck 4352 (USF
297911); Tampa, Learning Gate Community grounds, 4 May 2019, A. Farid 920 (epitype here
designated USF 301499); Palm Beach Co., Frenchman’s Reserve, 1 III 2019, J. Bolin 324 (USF
300081); Prosperity Oaks, 2 Mar 2019, J. Bolin 325 (USF 300080). OHIO: Hocking Co., 4 Aug
2018, J. Bolin 185 (USF 300091); Vinton Co. 5 Aug 2018, J. Bolin 184 (USF 300085).
TENNESSEE: Knox Co., Knoxville, Tobler Rd., 4 Sept 1949, A.J. Sharps s.n. with L.R. Hesler
(FLAS-F-53755).
Figure 4 – Field photographs of Cyanoboletus cyaneitinctus. A A. Farid 920. B A. Farid 340.
Macrochemical tests of basidiomes are labelled. The scale in the top is in centimeters. C JAB 324.
D JAB 389. Photo credits: J.A. Bolin.
Notes – Cyanoboletus Gelardi, Vizzini, & Simonini is in the Pulveroboletus clade, and is
comprised of eight species. Cyanoboletus was described in 2014 (Vizzini 2014) with Boletus
pulverulentus Opat. as the type species for the genus. Although no molecular analysis was provided
in the protologue, previous molecular analyses demonstrated several species (now in Cyanoboletus)

14
were not related to Boletus L. s. str. (Gelardi et al. 2013, Wu et al. 2014). Cyanoboletus is
distinguished from other boletoid genera by its yellowish brown to dark brown pileus, rapidly blueing
context and hymenophore, and smooth basidiospores.
Cyanoboletus cyaneitinctus is very similar to the closely related C. pulverulentus (Opat.)
Gelardi, Vizzini & Simonini. Both are boletes with a dark brown pileus, small pores (1–2 per mm),
and yellow stipes with brown punctae; all surfaces instantly bruise blue. The European name has
historically been applied to this species in North America (Singer 1947, Smith & Thiers 1971,
Bessette et al. 2000, 2017), but we are here treating them as separate species based on our molecular
analyses (Fig. 1) and morphological studies. The spore quotient Q is lower in C. cyaneitinctus at Q
= 2.4 (with the Q usually between 2.3–2.5) compared to 2.6–2.9 in C. pulverulentus (Gelardi et al.
2013). These two species are geographically separated, with C. cyaneitinctus occurring in eastern
North America and the latter found in Europe. Cyanoboletus sinopulverulentus, which is sister to
C. cyaneitinctus (Fig. 1) is distinguished from C. cyaneitinctus and C. pulverulentus by its evenly
dark brown stipe (lacking the reddish and yellow tones often present in the other two species), which
is more heavily pruinose to scissurate. Cyanoboletus sinopulverulentus has predominately 2-spored
basidia (4-spored in the other two taxa), and can also be distinguished on the basis of its Q value,
which is reported as 2.17–2.45 (Gelardi et al. 2013), smaller than either of the other two Cyanoboletus
species mentioned here.
Boletus mutabilis Morgan is an earlier but illegitimate name for this American species (see Art.
53.1). Thus, the oldest name we have to apply to the North American species is C. cyaneitinctus.
Singer (1947) treated C. cyaneitinctus as a synonym of C. pulverulentus. The type of C. cyaneitinctus
was examined, and matched the other North American collections examined. This type material is
quite old and not in good condition; thus, we have designated an epitype, and have included images
(Figs 4, 5A–G) as well as published molecular data. Cyanoboletus cyaneitinctus and C. pulverulentus
are difficult to distinguish morphologically.
Cyanoboletus cyaneitinctus forma reticulatus (Snell, E.A. Dick & Hesler) A. Farid comb. nov.
Fig. 5H
MycoBank number: MB 840859; Facesoffungi number: FoF 10465
Basionym – Boletus pulverulentus f. reticulatus Snell, E.A. Dick & Hesler, Mycologia 43(3):
362. 1951.
Typification – USA, Tennesse, Knox County, Knoxville, on an old sod yard near Robinia and
Ligustrum and not far from Ulmus but with no accurate indication of mycorrhizal associate, 4 Sept
1949, L.R. Hesler 19314 (holotype TENN-F-019314, isotype SFSU -F-000439).
Material examined – USA, Florida, Hillsborough Co., Brandon, under Quercus laurifolia, 5
Jun 2020, Farid 1035 (USF 301501).
Notes – Cyanoboletus cyaneitinctus f. reticulatus differs from the type form by the reticulation
present over the upper stipe. The protologue states all other macro- and micromorphological
characters are consistent, and this is consistent with our observations.
Hemileccinum floridanum J.A. Bolin, A.E. Bessette, A.R. Bessette, L.V. Kudzma, A. Farid & J.L.
Frank sp. nov. Figs 6, 10J–L
MycoBank number: MB 840861; Facesoffungi number: FoF 10464
Etymology – A reference to Florida where this species was first collected and described.
Typification – USA, Florida, Lake County, Lake Louisa State Park, 4 Sep 2016, J. A. Bolin
142 (holotype USF 301495).
Diagnosis – Medium-sized to large basidiocarps with a dry to slightly tacky, reddish brown to
chestnut brown pileus and a whitish stipe that becomes pale yellow at the apex and has a white basal
mycelium. The hymenophore is bright yellow when young, becomes darker brownish yellow as it
matures, and does not stain when bruised. It has white context that slowly stains yellow often from
the margin toward the center. The basidiospores measure (10-)13-16(-17) × 4.5-6 µm and are
elliptical. It fruits on the ground with oak from late spring through fall.

15
Figure 5 – Field photographs of Cyanoboletus cyaneitinctus. A JAB 325. B JAB 185. C–G JAB 324.
H Cyanoboletus cyaneitinctus f. reticulatus Farid 1035. Photo credits: A–G J.A. Bolin, H A Farid.
Description – Pileus 2.8-12.5 cm wide, convex becoming broadly convex to nearly plane in
age; surface dry to slightly tacky, smooth to somewhat wrinkled and uneven, glabrous to finely
velvety, sometimes with a whitish bloom when young, reddish brown to chestnut brown, cuticle
acidic tasting or not distinctive; margin even or nearly so. Hymenophore tubulose 3-12 mm deep,
pore surface bright yellow when young, maturing to darker brownish-yellow, not staining when
bruised, depressed near the stipe in age, easily detached from the pileus context; pores angular to
irregular, 2–3 per mm. Stipe 4-9.5 cm long × 1-3 cm thick, nearly equal or enlarged in either direction,
with a pinched base; surface dry, longitudinally striate, nearly glabrous to very weakly scurfy-
punctate, not reticulate; whitish overall on young specimens, becoming pale yellow at the apex with
variable reddish tints and streaks over a whitish to pale yellow ground color below, with white basal
mycelium. Context in the pileus white, slowly staining yellow often from the margin toward the
center, with a slight pinkish-red coloration beneath the cuticle; in the stipe, white, slowly staining
yellowish from the pileus trama just above the hymenophore partly downward along the exterior
stipe trama when exposed. Cuticle stains brownish red or light orange sometimes fading to light green
with the application of KOH, olive and then orange or amber with a green ring with NH4OH, and

16
dark orange-amber to orange with FeSO4. The context stains pale orange to yellow then fades with
KOH, is negative with NH4OH, and negative or light greyish olive green with FeSO4. Odor slightly
sour to not distinctive; taste not distinctive.
Basidiospores olive-brown in fresh deposit, (10–)13–16(–17) × 4.5–6 µm, n = 30, Q = 2.86,
elliptical in face view, inequilateral in profile, thin-walled, smooth, lacking an apical pore, grayish
yellow in KOH, brownish yellow in Melzer’s. Basidia 32-38 × 8.5-10.5 µm, clavate, 4-sterigmate,
sometimes 3- or 2-sterigmate, hyaline in KOH, yellow in Melzer’s, with granular, inamyloid
contents. Basidioles 22–29 × 7.5–8.5 µm, clavate, hyaline in KOH, yellow in Melzer’s.
Pleurocystidia 25–50 × 6–10 µm, hyaline, ventricose in the middle, ampullaceous at the apex,
frequent near the pores. Pileipellis a cutis of loosely interwoven cylindric hyphae with markedly
inflated, sphaerocyst-like oval to subglobose terminal cells, 11–33 × 15–22 µm, grayish yellow in
KOH, yellow to orange-yellow with granular contents in Melzer’s; hyphae of the pileipellis 4-8.5 µm
wide, thin-walled, smooth, grayish yellow in KOH, yellow in Melzer’s. Pileus trama hyphae loosely
interwoven, highly variable, 4–16 µm, with rounded terminal ends, thin-walled, smooth, hyaline in
KOH, yellow in Melzer’s, inamyloid. Hymenophoral trama boletoid, with lateral elements 5–9 µm,
moderately divergent, thin-walled, smooth, hyaline to pale grayish yellow in KOH, pale grayish
yellow in Melzer’s. Stipitipellis 4–17 µm wide, hyphae subparallel, highly variable, tubular with
rounded ends and granular contents, thin-walled, smooth, hyaline in KOH, yellow-brown in
Melzer’s, caulocystidia not observed. Stipe trama interwoven, 9–27 µm, highly variable, tubular with
rounded ends, thin-walled, smooth, hyaline in KOH, hyaline to pale yellow in Melzer’s. Clamp
connections absent.
Figure 6 – Field photographs of Hemileccinum floridanum. A J.A. Bolin 142. B. J.A. Bolin 454.
D J.A. Bolin 157. E J.A. Bolin 201. Photo credits: J.A. Bolin. C, F are SEM images of basidiospores
from J.A. Bolin 454 (white bar = 4 µm).
Habit, Habitat and Distribution – Solitary, scattered or in groups on the ground with oak; known
from Florida, potentially to North Carolina (Singer 1947).
Material examined – USA, Florida, Hillsborough County, Violet Cury Nature Preserve, 14 June
2017, A. Farid 625 (USF 301503); University of South Florida Tampa campus, trails near tennis
courts in NE corner of campus, 4 Jun 2018, A. Farid 790 (USF 297572), ibid., 30 Oct 2019, A. Farid

17
1032 (USF 301509). Lake County, Lake Louisa State Park, 4 Sept 2017, J.A. Bolin 142; Palm Beach
County, Frenchman’s Forest Natural Area, Jupiter 7 Nov 20 May 2018, J.A. Bolin 157 (USF
301488), ibid., 7 Nov 2019, J.A. Bolin 142 (USF 301495); ibid., 7 Nov 2019, J.A. Bolin 454 (USF
301491).
Notes – Hemileccinum subglabripes (Peck) R. Halling is very similar, but its stipe is
furfuraceous to scabrous or fibrillose and pale to bright yellow. It has smaller narrower spores and
smaller sphaerocyst-like elements that measure 10–24 µm. Hemileccinum hortonii (A.H. Sm. &
Thiers) M. Kuo & B. Ortiz is also similar, but it has a conspicuously pitted pileus, a smooth to lightly
pruinose stipe that sometimes has delicate reticulation on the upper half, and pores that sometimes
bruise blue. Hemileccinum rubropunctum (Peck) R. Halling & B. Ortiz has a conspicuously punctate
stipe, yellowish context, sometimes with an unpleasant odor reminiscent of stale cigarette butts in an
ashtray and larger spores that measure (10) 16–22 × 5.5–7.5 µm. The basidiospores under SEM were
smooth, lacking the very minute warts present in some species of Hemileccinum (Šutara 2008, Wu
et al. 2014).
Lanmaoa sublurida (Murrill) A. Farid & A.R. Franck comb. nov. Figs 7, 8, 10M–O
MycoBank number: MB 840862; Facesoffungi number: FoF 10463
Basionym – Suillellus subluridus Murrill, Mycologia 30(5): 524 (1938).
Typification – USA. FLORIDA: Alachua Co., Gainesville, Murrill (holotype FLAS-F-15869).
Synonyms – Boletus miniato-olivaceus var. subluridus (Murrill) Singer, Mycologia 37(6): 798
(1945); Boletus subluridus (Murrill) Murrill, Mycologia 30(5): 525 (1938).
Description – Pileus 3–14 cm wide, pulvinate when young, margin entire or wrinkled,
becoming convex to nearly plane at maturity, smooth, somewhat tacky when wet, bright red to ruby
red when young, becoming mixed with various shades of bright red, orange red, or a peach-colored
orange, especially at the margins, or sometimes turning brown entirely at maturity, sometimes
becoming rimulose, revealing the context color beneath the cuticle. Hymenophore tubulose, 5–12
mm deep at maturity, tubes sulphur yellow, bluing on injury, fading to olive green; pores initially
appearing yellow when young, stuffed, slowly and unevenly maturing to reveal red pore mouths at
maturity. Stipe 5–8 × 1.5–3 cm, equal, tapering upwards, or sometimes ventricose, pale yellow,
especially when young, with a small network of reticulation forming isodiametric meshes in a narrow
zone to 2 (–5) mm long at the apex of the stipe, but sometimes absent, especially in younger
specimens, and the rest of the stipe glabrous when young with fine floccons which develop over the
stipe surface as it matures, appearing smooth without a hand lens or without close inspection, at
maturity these floccons giving the appearance of a stipe that is red to purplish-red, stipe surface
bruising a light blue, especially when young; basal mycelium white to pale yellow. Context yellow
throughout, or sometimes yellow only in the stipital context (especially so when mature), not bluing
or only very weakly and slowly bluing when young, mature specimens bluing in the stipital and pileal
context around the hymenophore. Taste mild, odor disagreeable, fetid, ammonia-like and slightly
alliaceous.
Basidiospores (8.7)9.3–10.8(12.6) × (3)3.4–4(4.6) μm, n = 71, Q = 2.8, boletoid thick-walled,
ellipsoid-oblong to subcylindric or subfusoid, smooth, melleous. Basidia 20–25 × 6–8 μm, 2- or 4-
spored, thin-walled, hyaline, clavate to pyriform; sterigmata 1–2 μm, occasionally pigmented like
pluerocystidia. Basidioles similarly sized and shaped. Pleurocystidia 30–35 × 10–15 μm, pigmented
a light golden brown in KOH, NH4OH, H2O, and Melzer’s, spores generally are clustered onto
cystidia. Cheilocystidia 15–50 × 5–10 μm, moderately thin-walled (0.5 μm), usually pigmented like
the pleurocystidia, but occasionally hyaline. Hymenophoral trama divergent. Pileipellis elements
septate, terminal elements 20–65 × 4–10 μm, thin-walled, cylindrical, with filiform apices that are
occasionally clavate, forming an ixotrichodermium of erect elements, occasionally becoming
prostrate and forming an ixosubcutis. Pileal trama composed of interwoven hyphae 3–10 μm wide,
thin-walled, cylindrical.
Material examined – USA, Florida, Alachua County, Gainesville, Beneath Laurel Oak
[Quercus laurifolia], 3 Jul 2020, A. Farid 1058 with R.E. O’Donovan and C. Peyer (USF 301505).

18
Hillsborough County, Brandon, S of Camden Visconti pond at main entrance, 19 June 2016, A. Farid
343 (USF 288426); ibid., 26 Jun 2017, A. Farid 631 (USF 301506); ibid., 11 Jun 2018, A. Farid 805
(USF 298026); 22 Oct 2019; ibid., 11 Jun; A. Farid 1023 (USF 300104); Lithia, beneath Quercus
laurifolia, 15 Jul 2020, A. Farid 1072 (USF 301508); ibid., 16 Jul 2020, A. Farid 1073 (USF 301504).
Notes – Lanmaoa G. Wu & Zhu L. Zang is a genus of boletes which is typically distinguished
by its thin hymenophore (1/3–1/5 the thickness of the pileal context at a position halfway to the pileal
center), which stains blue when bruised, a light-yellow context which stains pale blue slowly when
cut, and an interwoven trichodermium to subcutis pileipellis. Although no molecular diagnosis was
provided in the paper describing the genus (Wu et al. 2015), the phylogenetic placement was based
on the work by Wu et al. (2014). Chai et al. (2018) describe the overlapping features of Lanmaoa
rubriceps N.K. Zeng & Hui Chai with Cyanoboletus, including hymenophore size, and staining
features. Lanmaoa sublurida is distinguished from similar looking boletes by the combination of its
characteristic odor, a pileus that varies in reds and orange that matures to a peach-orange, sometimes
brown, yellow tubes with pores that appear yellow and mature to carmine, and a light-yellow stipe
with fine floccons that densely cover it at maturity.
Figure 7 – Field photographs of Lanmaoa sublurida. A. Farid 1072. B. Farid 343. C. Farid 1073.
Photo credits: A Farid.

19
Figure 8 – Field photographs of Lanmaoa sublurida. A Farid 1072. B–C Farid 1078. D Farid 631.
E–F Farid 1072, cross-section of younger (E) and older (F) specimens. Photo credits: A. Farid.
There are several species in the southeastern USA that might be confused with L. sublurida.
Boletus carminiporus Bessette, Both & Dunaway, described from Mississippi, could be confused
with L. sublurida, although B. carminiporus differs in the lack of staining in the context at any stage,
lacks any distinctive odor, and its stipe is usually redder, and stains olive-brown, olive-green, to olive-
yellow. Lanmaoa borealis (A.H. Sm. & Thiers) A.E. Bessette, M.E. Nuhn & R.E. Halling is similar,
but has larger spores (11–13[15] µm long) and has only been documented from the northeastern
USA. The similar Boletus sensibilis Peck, found in the eastern USA, bruises similarly on the stipe,
but the pore mouths are yellow (never red), the stipe develops a flush of red on the bottom half (never
the purplish red that L. sublurida develops at maturity), and the context blues more readily
throughout.
Xerocomellus bolinii J.A. Bolin, A.E. Bessette, A.R. Bessette, L.V. Kudzma, J.L. Frank & A. Farid,
sp. nov. Figs 9, 10P–R
MycoBank number: MB 840863; Facesoffungi number: FoF 10462
Typification – USA, Florida: Broward County, Davie, Tree Tops Park, 27 Jan 2017, J.A. Bolin
43 (holotype USF 301496).
Etymology – The epithet bolinii honors Franklin Alexander Bolin, a biologist, naturalist and
educator who for more than twenty-five years introduced thousands of students to the fields of
mycology, herpetology and lepidoptery. Franklin was born and raised in Northeastern Ohio and
attended Ohio State University where he earned a master’s degree in both Field Zoology and
Herpetology. He went on to become an Advanced Biology teacher at Grove City High School from
1963–1988. Using his unique and progressive classroom style which immersed students in “hands
on learning”, Bolin developed a curriculum for the entire school district known as “The Natural
History of Ohio”.
Diagnosis – Small to medium-sized basidiocarps with a dry, blue-staining, appressed-fibrillose
to squamulose pileus with pinkish brown fibrils with white to creamy white context visible in the

20
cracks. The cap context is creamy white or a mixture of creamy white and yellow, becoming yellow
to orange in the stipe and rapidly stains blue when exposed. The pore surface is yellow when young,
becomes dull yellow at maturity, and rapidly stains blue when bruised. The stipe has reddish brown
punctae over a whitish to pale yellow ground color that darkens toward the base and staining blues
when bruised. Basidiospores measure (10–)12–13(–14) × 4.5–6 µm. It fruits on the ground with
Quercus and Pinus.
Description – Pileus 4-8 cm wide, convex becoming broadly convex to nearly plane in age;
pileus appressed-fibrillose to squamulose with pinkish brown fibrils and white to creamy white
context visible in the cracks, dry, staining blue, sometimes slowly or weakly; margin incurved at first
remaining so well into maturity, sterile, sometimes undulating, becoming conspicuously cracked with
age. Hymenophore tubulose, pale yellow, becoming dull yellow in age, quickly staining dark blue;
pores 1–2 per mm, angular to irregular or slightly elongated; tubes 2-6 mm deep, rapidly staining
blue when exposed. Stipe: 50–90 × 8–15 mm wide, nearly equal or slightly tapered downward, with
a pinched based, solid; surface dry, weakly longitudinally striate, with reddish brown punctae over a
whitish to pale yellow ground color often with reddish tints extending from the base upward, staining
blue when bruised, basal mycelium white to creamy white. Context of pileus creamy white or a
mixture of creamy white and yellow becoming yellow to orange in the stipe and rapidly stains blue
when exposed. Odor and taste not distinctive. Macrochemical Testing: Pileus of mature specimens
showed light green fading to yellow with NH4OH; younger specimens turn orange with faint green
outline of stained area, eventually fading to yellow. Orange to amber, fading to brown with KOH.
Older specimens light brown and younger specimens light green with FeSO4. Context in both mature
and younger specimens pale orange to NH4OH, orange to amber fading to brown with the application
of KOH and yellow with FeSO4. Cuticle stains light green, fading to yellow with NH4OH; younger
specimens develop orange with a faint green outline that eventually fades to yellow; KOH produces
orange to amber that fades to brown; with FeS04 mature specimens turn light brown, and younger
specimens light green.
Figure 9 – Field photographs of Xerocomellus bolinii. A J.A. Bolin 238. B J.A. Bolin 274.
C J.A. Bolin 232. D J.A. Bolin 208. Photo credits: J.A. Bolin.

21
Figure 10 – Microscopic structures of the boletes from this study. A–C Cyanoboletus cyaneitinctus.
D–F Aureoboletus pseudoauriporus. G–I Cyanoboletus bessettei. J–L Hemileccinum floridanum.
M–O Lanmaoa sublurida. P–R Xerocomellus bolinii. A, D, G, J, M, P depict the pileipellis for each
species (black bar = 50 µm), D showing a gelatinized pileipellis, B, E, H, K, N, Q depict basidiospores
with guttules (black bar = 10 µm), and C, F, I, L, O, R depict basidia, basidioles, and cystidia (black
bar = 20 µm), with guttules present. Drawing credits: A. Farid.
Basidiospores light brown to olive-brown in fresh deposit, (10–)12–13(–14) × 4.5–6 µm, n =
30, Q = 2.40, elliptical in face view, obscurely inequilateral in profile, thin-walled, smooth, lacking
an apical pore, pale grayish yellow in KOH, dull yellow in Melzer’s. Basidia 32-36 × 9-12 µm,
mostly clavate, occasionally cylindro-clavate, 2(4)-sterigmate, hyaline in KOH, grayish yellow in
Melzer’s. Basidioles 21.5–30 × 6.5–10 µm, clavate. Hymenophoral trama boletoid, with lateral
elements, 5-8 µm wide, moderately divergent, hyaline in KOH, grayish yellow in Melzer’s.
Pileipellis a tangled layer or repent tubular hyphae, 5–9.5 µm wide, with cylindrical, rounded to
slightly inflated end cells, thin-walled, smooth, hyaline in KOH, yellowish in Melzer’s. Pileus trama
hyphae loosely interwoven, 5–11 µm wide, smooth, thin-walled, hyaline in KOH or Melzer’s.
Cheilocystidia and pleurocystidia scattered, 36–48.5 × 9–11.5 µm, fusoid-ventricose, smooth, thin-
walled, hyaline to ochraceous in KOH, ochraceous in Melzer’s. Stipitipellis hyphae mostly parallel,
slightly interwoven, 4.5–9.5 µm wide, hyaline to yellowish in KOH or Melzer’s, with fascicles of
clavate to distorted caulocystidia 34–52 × 9–21 µm, that are dull yellow to brownish yellow in KOH
or Melzer’s. Stipe trama subparallel, interwoven, 5–11.5 µm wide, hyaline to yellowish in KOH or
Melzer’s. Clamp connections absent.

22
Habit, Habitat and Distribution – Solitary or scattered in sandy soil associated with Quercus
and Pinus, along woodland edges, typically near saw palmetto (Serenoa repens) and/or cabbage palm
(Sabal palmetto). Currently only documented from Florida. There are several images that we believe
to be X. bolinii on the citizen science platform MushroomObserver.org (observation nos.: 430943,
412138, 293427, 289394), but no herbarium samples were made.
Specimens examined – USA, Florida, Broward County, Davie, Tree Tops Park, 14 Oct 2017,
J.A. Bolin 124 (USF holotype 301494); Lake County, Lake Louisa State Park, Clermont, 13 Jun
2020, A. Farid 1047 with R.E. O’Donovan, C. Matson, and J.A. Bolin (USF 301498); Palm Beach
County, Delray Beach, Morikami Museum and Japanese Gardens, 23 Sep 2017, Jason Bolin 110
(USF 300098); ibid., 17 Oct 2018, J.A. Bolin 232 (USF 300082); ibid., 12 Sep 2018, J.A. Bolin 208
(USF 301486); West Palm Beach, Okeeheelee Park, 20 Nov 2017, J.A. Bolin 133 (USF 300094);
ibid., 13 Nov 2018, J.A. Bolin 238 (USF 301485); ibid., 23 Nov 2018, J.A. Bolin 274 (USF 301484).
Notes – Xerocomellus chrysenteron is similar but has a dark olive to olive-brown or greyish-
brown cracked cap with exposed red to pinkish context, stains slowly or erratically greenish-blue on
the hymenophore and cap context and has a more northern distribution. Xerocomellus zelleri has a
dull black to blackish-brown or dark olive-brown pileus, context that is white to pale yellow that is
unchanging or sometimes bluing and is reported from the Pacific Northwest south to California and
into Mexico.
Discussion
Contextualizing the species treated in this study
The species treated in this paper further our understanding of the boletes, both in terms of
biodiversity and systematics. Our analyses (Fig. 1) of Aureoboletus pseudoauriporus and its allies
indicate that A. auriporus (Peck) Pouzar represents a species complex. Aureoboletus auriporus was
described from New York as Boletus auriporus Peck (Peck 1873), with the protologue indicating a
grayish-brown, sometimes tinged with red pileus color. The name has been applied widely to
specimens across eastern North America, but specimens sequenced from Florida differ
phylogenetically from specimens in the northeast. The pileus in A. pseudoauriporus is pinkish-tan,
which differs from the grayish colors described in the protologue of Boletus auriporus. Our
phylogenetic analyses placed A. pseudoauriporus sister to a clade containing two specimens of A.
auriporus from Indiana and Tennesssee and A. viridiflavus from North Carolina. Three other
specimens of A. auriporus (from Massachusetts, North Carolina, and Costa Rica) fell separately
outside of this group (see Fig. 1). Aureoboletus viridiflavus, described from North Carolina, is a rarely
documented species that is often confused for A. auriporus. In a monograph of Aureoboletus, Klofac
(2010) noted most authors took A. viridiflavus as A. auriporus, but noted the subtle morphological
characters separating the two species.
We have expanded our understanding of North American Cyanoboletus with the resurrection
of C. cyaneitinctus as well as the addition of the novel species C. bessettei. Cyanoboletus
cyaneitinctus is widely distributed across North America. Many previous works on North American
boletes applied the European name C. pulverulentus to the North American species (Singer 1947,
Bessette et al. 2017). Phylogenies consistently show significant divergence between specimens from
North America and those from Europe (Gelardi et al. 2013, 2015, Braeuer et al. 2018, Fig. 1),
supporting the recognition of North American material as the species C. cyaneitinctus. So far, the
only other Cyanoboletus species known from North America now includes C. bessettei.
Cyanoboletus bessettei is only known from the type location in South Carolina, but we expect future
studies will better establish its geographical limits. Although briefly treated as a Cyanoboletus in a
study by Vizzini (2014), molecular analyses by Frank et al. (2020) have since shown Xerocomellus
rainisiae (Bessette & O.K. Mill.) N. Siegel, C.F. Schwarz & J.L. Frank is not a member of
Cyanoboletus. Morphological characters that X. rainisiae shares with Xerocomellus Šutara include
the pileus that becomes rimose in age, the deep red pigmentation of the basal stipital context (though
less extensive than typical Xerocomellus), and the bright yellow, blue-staining hymenium.

23
Similar to Chai et al. 2018, we have found Lanmaoa and Cyanoboletus to be closely related
(Fig. 1) and morphologically intergrading, although Cyanoboletus tends to have dull brown colors
and Lanmaoa often has bright red or yellow coloration (Wu et al. 2014, 2016, Chai et al. 2018).
Cyanoboletus bessettei and C. instabilis both share the 1/3–1/5 hymenophore-to-pileal-context ratio
found in Lanmaoa (and some Baorangia). Chai et al. (2018) suggested future research may consider
treating Cyanoboletus and Lanmaoa as one genus, in which Cyanoboletus would have priority over
Lanmaoa (Art. 11.3 of the Shenzhen Code). This is complicated by Vadthanarat et al.’s (2019)
phylogenetic inference of the genus Cacaoporus Raspé & Vadthanarat, which used the loci TEF1,
RPB2, atp6, and cox3 to place two named and one unnamed species of Cacaoporus sister to
Cyanoboletus, while receiving no phylogenetic support for Lanmaoa and Cyanoboletus as sister
genera. Due to the limited overlap of data between our dataset and Vadthanarat et al.’s dataset,
sequences of Cacaoporus were not included in our final analyses. We believe the suggestion by Chai
et al. (2018) to lump Lanmaoa and Cyanoboletus should be carefully re-considered in future studies
of this clade as more data become available.
Hemileccinum floridanum forms a well-supported sister clade to Hemileccinum subglrabipes,
the species it most closely resembles (Fig. 1). Using Singer (1947), Hemileccinum floridanum keys
out to Leccinum subglabripum (Peck) Sing. (= Hemileccinum subglabripes). Insightfully, under his
L. subglabripum, Singer (1947) gave a separate description for the Florida collections, which here
conform to the new species H. floridanum. Leccinum subglapripes var. corrugatoides Singer was
also described in Singer (1947), but differs from H. floridanum by a very rugose, “light brownish
olive” pileus and a “light brownish olive” spore print (Singer 1947). Our collections do not possess
these features, and it remains to be determined if this taxon is distinct from H. floridanum. Molecular
analyses by Kuo & Ortiz-Santana (2020) revised the concept of Hemileccinum to include H.
rubropunctum, a widespread species in North America which forms tuberculate ectomycorrhizae
with Quercus spp. (Smith & Pfister 2009). Roots beneath several collections of Hemileccinum
floridanum were examined for tuberculate ectomycorrhizae, but none were located. Thus far H.
rubropunctum is unique in its ability to form tuberculate ectomycorrhizae within the Boletaceae,
though other Boletales are capable of this (e.g. Suillus and Rhizopogon).
Xerocomellus bolinii is here placed as sister to a clade of Eurasian species, one of which has
been considered part of the genus Rheubarbariboletus Vizzini, Simonini & Gelardi. Similar to Frank
et al. (2020), our phylogenetic analysis finds Heliogaster Orihara & K. Iwase and
Rheubarbariboletus embedded in the Xerocomellus lineage and Nigroboletus to be sister to this
broadly defined Xerocomellus lineage. Vizzini (2015) cited the ITS-based phylogeny in Gelardi et
al. (2013) and unpublished data as the molecular basis for establishing Rheubarbariboletus, differing
from Xerocomellus by its smooth, non-striate and non-truncate spores, smooth or finely incrusted
pileipellis, congophilous plaques on the hyphal surface, tapered and rooting stipe base, the bright
yellow-ochraceous to orange-rhubarb and unchangeable context in the stipe base, and the dark blue-
green blackish reaction with FeSO4 on the pileus surface and the base of the stipe context.
Xerocomellus bolinii, while sharing the non-bluing basal stipital context, smooth, non-truncate
spores, and smooth pileipellis of Rheubarbariboletus, differs in its reaction to the application of
FeSO4 to the context by only turning light brown to light green (in old and young specimens,
respectively), and by lacking a rooting stipe. In light of these molecular and morphological data, it
seems best to include Heliogaster and Rheubarbariboletus within Xerocomellus at this time.
We follow Bozok et al. (2019) in recognizing Exsudoporus as a genus separate from
Butyriboletus. Wu et al. (2016) treated the genus as a synonym of Butyriboletus, citing the reticulation
and interwoven trichodermium to subcutis pileipellis as shared characters with the genus
Butyriboletus. Bozok et al. (2019) reported on the positive amyloid reaction in the stipe tissues of E.
permagnificus, a feature not shared with Butyriboletus. Our observations of E. floridanus show that
the stipe base context exhibits a dextrinoid reaction in the stipe base (pers. obs.). Also, Exsudoporus
species have pores that are discolorous with the tubes, and the basidiomes bruise blue much darker
and heavier than Butyriboletus species. The guttation on the pores is regularly found, especially in
younger specimens of Exsudoporus species, and is a useful distinguishing character. Wu et al. (2016)

24
reported species which were sister to the clade containing Exsudoporus and Butyriboletus, but these
species remain undescribed. Additional analyses and thorough morphological comparison of those
undescribed species might justify a broader concept of Butyriboletus, however until those analyses
are produced, retaining the genus Exsudoporus is preferred.
Conclusions
This paper updates our understanding of the boletes in southeastern USA. Four novel species
are described, as well as resurrecting and applying the name Cyanoboletus cyaneitinctus to the
Cyanoboletus species widespread across North America. Our molecular analyses (Fig. 1) provide a
DNA-based approach to aid morphological classification of these boletes and to better understand
the distribution of these species. Our analyses also support the many genera found in recent
Boletaceae phylogenetic reconstructions (Wu et al. 2014, 2016). By analyzing the protein-coding loci
(RPB1, RPB2, TEF1) from a collection of Butyriboletus floridanus on GenBank, we have also
confirmed a disjunct distribution for this tropical species. Inclusion of additional data from the
epitype of Pulchroboletus rubricitrinus also lends the specimen to broader phylogenetic analyses.
We also provide sequences of western USA Xerocomellus, which will aid future bolete phylogenetic
reconstructions, as many species of Xerocomellus from North America lack protein coding loci.
This paper increases the knowledge of biodiversity present in the region. The potential for
robust future studies is impeded by a lack of baseline knowledge of biodiversity. As molecular
phylogenetic analyses continuously update the taxonomy of our classifications of the boletes, the
need for further investigations into the boletes of the southeastern USA becomes readily apparent.
Important aspects, such as morphological traits, host-specificity and geographic distribution, have
been shown to be incredibly important with regards to boletes. Species-level concepts which were
once broadly defined and applied widely across eastern North America have been shown to
encompass several species, sometimes with clear morphological characters to distinguish them, as
well as cryptic species in which geography seems to play a key role. Increasing and updating our
understanding of boletes allows researchers to obtain richer species-level sequence-based
identifications in environmental studies (Hibbett et al. 2011, Truong et al. 2017, Xu 2016), which is
important for ecological studies, and paramount to better understanding threatened ecosystems in the
southeastern USA. Macrofungal species have shown the potential for introduction and spread, e.g.
Favolaschia, Clathrus archeri, Perenniporia ochroleuca, and the bolete Aureoboletus projectellus
(Desprez-Loustau et al. 2007, Pringle et al. 2009, Vizzini et al. 2009, Wrzosek et al. 2017, Banasiak
et al. 2019). Considering many species of boletes in the southeastern USA are geographically
restricted, there is the potential that exotic mycorrhizal fungi may outcompete these endemic species.
Acknowledgements
This work was funded by the Cooley and Lakela funds at USF. The authors would like to thank
the herbaria staff and faculty who assisted in facilitating loans for this study. We would also like to
thank Hillsborough County Conservation and Environmental Lands Management, the Southwest
Florida Water Management District, the Florida Park Service’s Bureau of Natural & Cultural
Resources, and the Everglades National Park for granting us research permits that enabled us to
sample fungi from their respective parks and are grateful to the park managers and staff for their
support and assistance. We would also like to thank Jonathan Frank for sending USF specimens of
western Xerocomellus for our phylogenetic analyses, and Clare Dennison for her help at the SEM lab
at USF.
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Zhao K, Wu G, Yang ZL. 2014 – A new genus, Rubroboletus, to accommodate Boletus sinicus and
its allies. Phytotaxa 188(2), 61–77.
Supplementary Table 1 GenBank accession number and other information of sequences used in
phylogenetic analyses in this study. Sequences in bold were generated in this study.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Alessioporus
ichnusanus
AMB 12756
KJ729491
KJ729504
–
–
KJ729513
Alessioporus
ichnusanus
MG420a
KJ729496
KJ729509
–
–
–
Alessioporus
rubriflavus
ARB1356
KU736957
MH656696
–
–
–
Alessioporus
rubriflavus
JLF2561
KU736958
KC812306
–
–
–
Aureoboletus
auriflammeus
CFMR BOS
699
–
MK601706
–
MK766269
MK721060
Aureoboletus
auriporus
35 97
–
DQ534636
–
–
–
Aureoboletus
auriporus
AB11
MH796985
–
–
–
–
Aureoboletus
auriporus
AB12
MH796989
–
–
–
–
Aureoboletus
auriporus
BDCR0431
–
HQ161871
HQ161840
–
–
Aureoboletus
auriporus
FLAS F 60185
MH796985
–
–
–
–
Aureoboletus
auriporus
FLAS F 60914
MH211684
–
–
–
–
Aureoboletus
auriporus
FLAS F 60985
MH016931
–
–
–
–
Aureoboletus
auriporus
MAC09 TENN
MF755267
–
–
–
–
Aureoboletus
auriporus
S D Russell
MycoMap
6611
MK560093
–
–
–
–
Aureoboletus
catenarius
GDGM45142
–
MN204514
–
–
–
Aureoboletus
catenarius
HKAS54463
–
KT990509
KT990890
KT990348
KT990710

29
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Aureoboletus
catenarius
HKAS54467
–
KT990510
–
KT990349
KT990711
Aureoboletus
clavatus
GDGM42962
–
KR052045
KR052056
–
–
Aureoboletus
clavatus
GDGM42963
–
KR052046
KR052057
–
KR052054
Aureoboletus
clavatus
GDGM42984
–
KR052047
–
–
KR052055
Aureoboletus
clavatus
HKAS59802
–
KR052044
–
–
KR052053
Aureoboletus
duplicatoporus
HKAS63009
–
KT990511
KT990891
KT990350
KT990712
Aureoboletus
duplicatoporus
HKAS83115
–
KT990512
KT990892
KT990351
KT990713
Aureoboletus
gentilis
ADK4865
–
–
–
KT823994
KT824027
Aureoboletus
gentilis
MG372a
–
KF112344
KF112557
KF112741
KF134014
Aureoboletus
gentilis
Pug1
–
DQ534635
–
–
KF030399
Aureoboletus
griseorufescens
GDGM28490
–
MH670278
–
MH700241
–
Aureoboletus
griseorufescens
ZM131
–
MH670279
MH700220
MH700242
–
Aureoboletus
innixus
CFMR BOS
544
–
MK601707
–
MK766270
MK721061
Aureoboletus
innixus
MB03 104
–
KF030239
–
–
KF030400
Aureoboletus
mirabilis
HKAS57776
–
KF112360
KF112624
KF112743
KF112229
Aureoboletus
mirabilis
REH9765
–
KP327661
–
–
KP327709
Aureoboletus
moravicus
MG374a
–
KF112421
KF112559
KF112745
KF112232
Aureoboletus
moravicus
VDKO1120
–
–
–
MG212615
MG212573
Aureoboletus
moravicus f luteus
PARMA 1544
11
KJ676960
KJ676958
–
–
KJ676959
Aureoboletus
nephrosporus
HKAS67931
–
KT990516
KT990895
KT990357
KT990720
Aureoboletus
nephrosporus
HKAS74929
–
KT990517
KT990896
KT990358
KT990721
Aureoboletus
projectellus
MICH KUO
09111014
–
MK601708
–
MK766271
MK721062
Aureoboletus
projectellus
NYBG13392
–
KP327622
–
–
KP327675
Aureoboletus
quercus
spinosae
GDGM43755
KY039954
KY039967
KY039963
KY039958
–
Aureoboletus
quercus
spinosae
GDGM43758
KY039955
KY039968
KY039964
KY039959
–
Aureoboletus
quercus
spinosae
GDGM43786
–
KY039969
KY039965
KY039960
–
Aureoboletus
raphanaceus
GDGM44832
–
MH670268
MH700218
MH700236
MH700194

30
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Aureoboletus
raphanaceus
GDGM52543
–
MH670271
–
MH700237
–
Aureoboletus
raphanaceus
GDGM52590
–
MH670272
MH700219
MH700238
MH700193
Aureoboletus
roxanae
CFMR BOS
698
–
MK601709
–
MK766272
MK721063
Aureoboletus
roxanae
DS626 07
–
KF030311
KF030381
–
KF030402
Aureoboletus
russellii
CFMR BOS
716
–
MK601710
–
MK766273
MK721064
Aureoboletus
singeri
CFMR BZ
2395 BOS 468
MN250221
MK601711
–
MK766274
MK721065
Aureoboletus sp
GDGM44829
–
KY039970
–
KY039961
–
Aureoboletus
tenuis
GDGM42601
KF265358
KF534789
–
KT291754
KT291745
Aureoboletus
tenuis
HKAS75104
–
KT990518
KT990897
KT990359
KT990722
Aureoboletus
thibetanus
HKAS57692
–
KT990524
KT990901
KT990365
KT990728
Aureoboletus
thibetanus
HKAS76655
–
KF112420
KF112626
KF112752
KF112236
Aureoboletus
thibetanus
HKAS89494
–
KT990525
KT990902
KT990366
KT990729
Aureoboletus
tomentosus
HKAS59694
–
KT990513
KT990893
KT990352
KT990714
Aureoboletus
tomentosus
HKAS80485
–
–
KT990894
KT990353
KT990715
Aureoboletus
viridiflavus
–
AY612805
–
–
–
–
Aureoboletus
viscidipes
HKAS77103
–
KT990519
–
KT990360
KT990723
Aureoboletus
viscosus
OR0361
–
–
–
MH614751
MH614703
Aureoboletus
yunnanensis
HKAS75050
–
KT990520
KT990898
KT990361
KT990724
Aureoboletus
zangii
HKAS74751
–
KT990521
KT990899
KT990362
KT990725
Aureoboletus
zangii
HKAS74766
–
KT990522
KT990900
KT990363
KT990726
Baorangia
alexandri
EE 2018a LE
254265
MH043612
MH036170
–
–
–
Baorangia
alexandri
EE 2018a LE
254266
MH043611
MH036169
–
–
–
Baorangia bicolor
MB07 001
–
KF030246
KF030370
–
KF030405
Baorangia
emileorum
GS 10213
MH043613
MH036171
–
–
–
Baorangia
emileorum
PRM 934960
MH043616
MH036174
–
–
–
Baorangia
emileorum
TO HG131114
MH043617
MH036175
–
–
–
Baorangia
emileorum
TO HG171015
MH043615
MH036173
–
–
–
Baorangia
emileorum
TO HG191015
MH043614
MH036172
–
–
–
Baorangia major
OR209
–
–
–
MG897441
MG897431

31
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Baorangia major
OR404
–
–
–
MG897442
MG897432
Baorangia major
OR486
–
–
–
MG897443
MG897433
Baorangia
pseudocalopus
HKAS75739
–
KJ184558
KJ184564
KM605179
KJ184570
Baorangia
rufomaculata
BOTH4414
–
KF030248
KF030369
MG897435
KF030406
Boletellus
longicollis
HKAS53398
–
KF112376
KF112625
KF112755
KF112238
Boletellus
projectellus
AFTOL ID 713
AY789082
AY684158
–
AY787218
AY879116
Boletellus singeri
VB4530
–
KP327669
–
–
KP327713
Boletus
abruptibulbus
4588
–
KF030302
KF030388
–
KF030401
Boletus aff
amygdalinus
HKAS57262
–
KF112316
KF112501
KF112660
KF112174
Boletus
albobrunnescens
REH8790
KF668279
HQ161879
HQ161877
–
–
Boletus
amygdalinus
112605ba
–
JQ326996
KF030360
–
JQ327024
Boletus
austroedulis
REH8969
JN020990
HQ161847
HQ161816
–
–
Boletus edulis
BD380
EU231984
HQ161848
HQ161817
–
–
Boletus edulis
Be3
–
KF030282
GU187444
GU187774
GU187682
Boletus edulis
HMJAU4637
–
KF112455
KF112586
KF112704
KF112202
Boletus edulis
Trudell 03 289
09
EU231983
EU232006
EU231999
–
–
Boletus rubriceps
Arora11331
KC900403
KC900404
–
–
–
Boletus rubriceps
MICH KUO
08150719
–
MK601722
–
MK766284
MK721076
Boletus
semigastroideus
PBM 3076
JX258840
KF030352
KF030384
–
KF030430
Boletus separans
DPL 2704
–
KF030329
KF030385
–
KF030431
Boletus separans
MICH KUO
06201002
–
MK601723
–
MK766285
MK721077
Buchwaldoboletus
lignicola
HKAS76674
–
KF112350
KF112642
KF112819
KF112277
Buchwaldoboletus
lignicola
HKAS84904
–
KT990538
–
KT990377
KT990740
Buchwaldoboletus
lignicola
Pul1
–
JQ326997
–
–
JQ327040
Buchwaldoboletus
lignicola
VDKO1140
–
–
–
MH614756
MH614710
Butyriboletus
appendiculatus
BR502008929
55
KJ605668
KJ605677
KJ619481
–
–
Butyriboletus
appendiculatus
BR502008933
90
KT002598
KT002609
KT002621
–
KT002633
Butyriboletus
appendiculatus
Bap1
–
AF456837
KF030359
–
JQ327025
Butyriboletus
appendiculatus
MB000286
KT002599
KT002610
KT002622
–
KT002634
Butyriboletus
brunneus
NY00013631
KT002600
KT002611
KT002623
–
KT002635
Butyriboletus
pseudoregius
BR502015335
59 51
KT002603
KT002614
KT002626
–
KT002638

32
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Butyriboletus
pseudoregius
BR502016184
65 02
KT002602
KT002613
KT002625
–
KT002637
Butyriboletus
pseudospeciosus
HKAS63513
–
KT990541
KT990909
KT990380
KT990743
Butyriboletus
pseudospeciosus
HKAS63596
–
KT990542
KT990910
KT990381
KT990744
Butyriboletus
pseudospeciosus
N K Zeng2127
MH885349
MH879687
–
–
MH879716
Butyriboletus
regius
11265
–
KF030267
–
–
KF030411
Butyriboletus
regius
MB 000287
KT002605
KT002616
KT002628
–
KT002640
Butyriboletus
roseoflavus
HKAS54099
KJ909519
KY418892
KF739741
KF739703
KF739779
Butyriboletus
roseoflavus
HKAS63593
KJ909517
KJ184559
–
–
KJ184571
Butyriboletus
roseoflavus
N K Zeng2123
MH885348
MH879686
–
–
MH885348
Butyriboletus
subsplendidus
HKAS50444
–
KT990540
KT990908
KT990379
KT990742
Butyriboletus
yicibus
HKAS57503
KT002608
KT002620
KT002632
–
KT002644
Butyriboletus
yicibus
HKAS68010
–
KT002619
KT002631
–
KT002643
Chalciporus aff
piperatus
HKAS50214
JQ928610
JQ928621
JQ928594
–
–
Chalciporus
piperatus
HKAS84882
–
KT990562
–
KT990397
KT990758
Chalciporus
pseudorubinellus
4302
–
KF030284
–
–
KF030441
Chalciporus
rubinelloides
HKAS57362
–
KT990563
–
KT990398
KT990759
Chalciporus
rubinelloides
HKAS58728
–
KT990564
–
KT990399
KT990760
Chalciporus
rubinelloides
HKAS74952
–
KT990565
–
KT990400
KT990761
Corneroboletus
indecorus
OR0863
–
–
–
MH614772
MH614726
Cyanoboletus
HKAS76850
–
KF112343
KF112527
KF112697
KF112187
Cyanoboletus
brunneoruber
HKAS80579 1
–
KT990568
KT990926
KT990401
–
Cyanoboletus
brunneoruber
HKAS80579 2
–
KT990569
KT990927
KT990764
–
Cyanoboletus
hymenoglutinosus
AB 2016
KT860060
–
–
–
–
Cyanoboletus
pulverulentus
18188
JF907794
–
–
–
–
Cyanoboletus
pulverulentus
A21
JX434686
–
–
–
–
Cyanoboletus
pulverulentus
A7
JX434685
–
–
–
–
Cyanoboletus
pulverulentus
ASIS22672
KP004920
–
–
–
–
Cyanoboletus
pulverulentus
B21 specimen
PRM 935923
LT714704
–
–
–
–

33
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Cyanoboletus
pulverulentus
B23 specimen
PRM 944014
LT714705
–
–
–
–
Cyanoboletus
pulverulentus
B24 specimen
PRM 944001
LT714706
–
–
–
–
Cyanoboletus
pulverulentus
B25 specimen
PRM 944013
LT714707
–
–
–
–
Cyanoboletus
pulverulentus
B26 specimen
PRM 944022
LT714708
–
–
–
–
Cyanoboletus
pulverulentus
B27 specimen
PRM 935997
LT714709
–
–
–
–
Cyanoboletus
pulverulentus
CA050916 04
HM347646
–
–
–
–
Cyanoboletus
pulverulentus
JMP0012
EU819453
–
–
–
–
Cyanoboletus
pulverulentus
MG 126a
KT157053
KT157062
–
–
–
Cyanoboletus
pulverulentus
MG 456a
KT157054
KT157063
–
–
–
Cyanoboletus
pulverulentus
MG 628a
KT157055
KT157064
–
KT157069
–
Cyanoboletus
pulverulentus
RT00004
EU819502
–
–
–
–
Cyanoboletus
pulverulentus
RW109
–
–
–
KT824013
–
Cyanoboletus
sinopulverulentus
HMAS 266894
KC579402
–
–
–
–
Cyanoboletus sp
B28
LT714710
MF373585
–
–
–
Cyaonoboletus
instabilis
HKAS 59554
–
KF112412
KF112528
KF112698
KF112186
Exsudoporus
floridanus
CFMR BZ
3170
MN250222
MK601725
–
MK766287
MK721079
Exsudoporus
frostii
TENN 067311
KT002601
KT002612
KT002624
–
KT002636
Gymnogaster
boletoides
NY01194009
–
KT990572
KT990928
KT990406
KT990768
Gymnogaster
boletoides
REH9455
–
JX889673
–
–
JX889683
Heimioporus
australis
REH9288
–
KP327652
–
–
KP327703
Heimioporus
conicus
N K Zeng3109
MH241052
MH241051
–
–
MH241053.
Heimioporus
cooloolae
REH9817
–
KP327664
–
–
KP327710
Heimioporus
cooloolae
REH9852
–
KP327665
–
–
KP327711
Heimioporus
gaojiaocong
N K Zeng2788
–
MF962380
–
–
MF962410
Heimioporus
gaojiaocong
N K Zeng2791
MF962398
MF962383
–
–
MF962412
Heimioporus
gaojiaocong
N K Zeng2792
MF962399
MF962384
–
–
MF962413
Heimioporus
gaojiaocong
N K Zeng2864
MF962400
MF962385
–
–
MF962415
Heimioporus
gaojiaocong
Z L Yang5901
MF962394
MF962377
–
–
MF962409

34
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Heimioporus
japonicus
HKAS52237
–
KF112347
KF112618
KF112806
KF112228
Heimioporus
japonicus
Lancang Y J
Hao84
MF962402
MF962386
–
–
MF962416
Heimioporus
japonicus
N K Zeng1335
MF962404
MF962388
–
–
MF962418
Heimioporus
japonicus
N K Zeng1566
–
MF962389
–
MF962424
MF962419
Heimioporus
japonicus
OR114
–
–
–
KT824004
KT824037
Heimioporus
subretisporus
HKAS80581
–
KT990573
–
KT990407
KT990769
Heimioporus
subretisporus
HKAS80582
–
KT990574
–
KT990409
KT990770
Hemileccinum
depilatum
AF2845
–
–
–
MG212633
MG212591
Hemileccinum
impolitum
Bim1
–
–
KF030375
–
JQ327034
Hemileccinum
impolitum
HKAS84869
–
KT990575
KT990930
KT990410
KT990771
Hemileccinum
rubropunctum
FH MES116
FJ480434
–
–
–
–
Hemileccinum
rubropunctum
FH MES117
FJ480433
–
–
–
–
Hemileccinum
rubropunctum
JLF5666
MH190826
MK874830
–
–
–
Hemileccinum
rubropunctum
NY01193924
–
MK601769
–
MK766328
MK721123
Hemileccinum
rubropunctum
NY792788
–
MK601768
–
MK766327
MK721122
Hemileccinum
rugosum
HKAS50284
–
KT990576
–
KT990411
KT990772
Hemileccinum
rugosum
HKAS84355
–
KT990578
KT990931
KT990413
KT990774
Hemileccinum
rugosum
HKAS84970
–
KT990577
–
KT990412
KT990773
Hemileccinum
subglabripes
MICH KUO
07070702
–
MK601737
–
MK766299
MK721091
Hemileccinum
subglabripes
MICH KUO
07230802
–
MK601738
–
MK766300
MK721092
Hemileccinum
subglabripes
MICH KUO
08301402
–
MK601739
–
MK766301
MK721093
Hemileccinum
subglabripes
MO 294169
MN128237
MN128238
–
–
–
Hemileccinum
subglabripum
72206
–
KF030303
KF030374
–
KF030404
Hortiboletus aff
rubellus
HKAS51239
–
KF112425
KF112618
KF112695
KF112184
Hortiboletus
amygdalinus
HKAS54166
–
KT990581
KT990933
KT990416
KT990777
Hortiboletus
amygdalinus
HKAS54242
–
KT990580
–
KT990415
KT990776
Hortiboletus
campestris
MICH KUO
08240502
–
MK601740
–
MK766302
MK721094

35
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Hortiboletus cf
rubellus
East Coast
MB03 033
–
–
KF030371
–
KF030419
Hortiboletus cf
rubellus
West Coast
PBM 1331
–
–
–
–
KF030420
Hortiboletus
indorubellus
DC 14
KT319647
KU566807
–
–
–
Hortiboletus
indorubellus
LS15
MK002767
MK002872
–
–
–
Hortiboletus
rubellus
MICH KUO
06081002
–
MK601741
–
MK766303
MK721095
Hortiboletus
rubellus
VDKO0403
–
–
–
MH614774
–
Hortiboletus
subpaludosus
HKAS52659
–
KT990582
–
KT990417
KT990778
Hortiboletus
subpaludosus
HKAS68158
–
KT990583
KT990934
KT990418
KT990779
Hymenogaster
behrii
OSC
Trappe12988
KJ882288
–
–
–
–
Hymenogaster
behrii
OSC
Trappe17620
KJ882290
–
–
–
–
Hymenogaster
macmurphyi
OSC MES282b
KJ882289
KJ882291
–
–
–
Lanmaoa
angustispora
HKAS74752
–
KM605139
KM605166
KM605177
KM605154
Lanmaoa
angustispora
HKAS74759
–
KM605140
KM605167
KM605178
KM605155
Lanmaoa asiatica
HKAS54095
–
KM605141
KM605164
KM605174
KM605151
Lanmaoa asiatica
HKAS63516
–
KT990584
KT990935
KT990419
KT990780
Lanmaoa asiatica
HKAS63592
–
KM605142
KM605163
KM605175
KM605152
Lanmaoa asiatica
HKAS63603
–
KM605143
KM605165
KM605176
KM605153
Lanmaoa asiatica
N K Zeng2125
MG030477
MG030470
–
–
MG030481
Lanmaoa asiatica
N K Zeng2795
–
MG030469
–
–
MG030480
Lanmaoa asiatica
OR0228
–
–
–
MH614777
MH614730
Lanmaoa borealis
2858
–
JQ326998
–
–
JQ327021
Lanmaoa
carminipes
MB06 061
–
JQ327001
KF030363
–
JQ327022
Lanmaoa cf
borealis AB35
MH796994
–
–
–
–
Lanmaoa
flavorubra
NY775777
–
JQ924339
–
KF112681
KF112160
Lanmaoa
macrocarpa
N K Zeng3021
–
–
–
–
MH879713
Lanmaoa
macrocarpa
N K Zeng3251
MH885347
MH879685
–
–
MH885347
Lanmaoa
pallidorosea
BOTH4432
–
–
–
MG897437
MG897427
Lanmaoa
pallidorosea
MO 210760
–
MH216001
–
–
MH318610
Lanmaoa
pallidorosea
MO 247881
MH234471
MH230088
–
–
MH337278
Lanmaoa
pseudosensibilis
DS615 07
–
KF030257
–
–
KF030407
Lanmaoa
roseocrispans
HOLOTYPE
–
MH036169
–
–
KP327616
Lanmaoa
rubriceps
N K Zeng2773
MG030475
MG030468
–
–
MG030479

36
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Lanmaoa
rubriceps
N K Zeng3006
MH885346
MH879683
–
–
MH879712
Nigroboletus
roseonigrescens
GDGM 43238
KT220584
KT220588
KT220591
–
KT220588
Nigroboletus
roseonigrescens
MG 524a
KT220586
KT220590
KT220593
–
–
Nigroboletus
roseonigrescens
ZT 13553
KT220585
KT220589
KT220592
KT220594
KT220596
Pulchroboletus
roseoalbidus
AMB 12757
KJ729486
NG_060126
–
–
KJ729512
Pulchroboletus
roseoalbidus
MCVE 17577
KJ729490
KJ729503
–
–
–
Pulchroboletus
roseoalbidus
MCVE 18217
KJ729488
KJ729501
–
–
–
Pulchroboletus
roseoalbidus
MG416a
KJ729489
KJ729502
–
–
–
Pulchroboletus
roseoalbidus
MG532a
KJ729487
KJ729500
–
–
–
Pulchroboletus
sclerotiorum
FLAS F 60333
MF098659
MF614166
MF614168
MF614169
MF614167
Pulchroboletus
sclerotiorum
FLAS F 60334
MF098660
–
–
MF614164
MF614165
Pulchroboletus
sclerotiorum
MO 243879
–
MH257545
–
–
MH337281
Pulveroboletus
auriporus
DD971
–
AY612819
–
–
–
Sinoboletus
duplicatoporus
HKAS50498
–
KF112361
KF112561
KF112754
KF112230
Suillellus
amygdalinus
NY00035656
–
KT990650
KT990990
KT990477
KT990840
Suillellus
amygdalinus
NY00815464
–
KT990659
KT990997
KT990484
KT990848
Suillellus queletii
VDKO1185
–
–
–
MH645604
MH645598
Suillellus
subamygdalinus
HKAS53641
–
KT990651
KT990991
KT990478
KT990841
Suillellus
subamygdalinus
HKAS57953
–
KT990652
KT990992
–
KT990842
Suillellus
subamygdalinus
HKAS74745
–
KT990653
KT990993
KT990479
KT990843
Xerocomellus
armeniacus
MA Fungi
47678
AJ419221
–
–
–
–
Xerocomellus
armeniacus
CM058
KP826760
–
–
–
–
Xerocomellus
armeniacus
ML41842RP
MH011927
–
–
–
–
“Rheubarbaribolet
us” persicolor
17602
JF908795
–
–
–
–
“Rheubarbaribolet
us” persicolor
SOMF 29860
MH011931
–
–
–
–
“Rheubarbaribolet
us” persicolor
SOMF 298154
MH011932
–
–
–
–
Xerocomellus
chrysenteron
HKAS56494
–
KF112357
KF112526
KF112685
KF112172
Xerocomellus
chrysenteron
MICH KUO
07271202
–
–
–
MK766373
MK721171

37
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Xerocomellus
chrysenteron
MICH KUO
09260903
–
–
–
MK766374
MK721172
Xerocomellus
chrysenteron
VDKO0821
–
–
–
KT824017
KT824050
Xerocomellus
chrysenteron
Xch1
–
–
KF030365
–
KF030415
Xerocomellus
cisalpinus
ADK4864
–
–
–
KT823993
KT824026
Xerocomellus
cisalpinus
AT2005034
–
–
KF030367
–
KF030417
Xerocomellus
cisalpinus
PDD94421
–
JQ924322
KF112525
KF112686
KF112171
Xerocomellus
communis
HKAS50467
–
KT990670
KT991008
KT990494
KT990858
Xerocomellus
communis
HKAS68204
–
–
KT991009
KT990495
KT991009
Xerocomellus
corneri
HKAS52503
–
KT990668
KT991006
KT990492
KT990856
Xerocomellus
corneri
HKAS90206
–
KT990669
KT991007
KT990493
KT990857
Xerocomellus
porosporus
VDKO0311
–
–
–
MH614773
MH614727
Xerocomellus
ripariellus
VDKO0404
–
–
–
MH614793
MH614746
Xerocomellus sp
HKAS50466
–
KF112372
KF112549
KT990494
KF112183
Xerocomellus sp
HKAS50467
–
KF112489
KT991008
KF112770
KF112173
Xerocomellus sp
HKAS51292
–
KF112369
KF112547
KF112692
KF112181
Xerocomellus sp
HKAS56311
–
KF112340
KF112524
KF112684
KF112170
Xerocomellus sp
HKAS59608
–
KF112371
KF112551
KF112696
KF112185
Xerocomellus sp
HKAS76673
–
KF112370
KF112548
KF112693
KF112182
Xerocomellus
zelleri
JLF2977
KM213666
KU144799
–
–
–
Xerocomellus
zelleri
REH8724
–
KF030271
KF030366
–
KF030416
Xerocomus
hortonii
MICH-KUO
07050706
–
MK601821
–
MK766377
MK721175
Xerocomellus
armeniacus
MA-Fungi
47678
AJ419221
–
–
–
–
Xerocomellus
persicolor
ML41842RP
MH011927
–
–
–
–
Boletus
pakistanicus
JQ178324
–
–
–
–
Bovista himalaica
JN411938
–
–
–
–
Xerocomellus
17602
JF908795
–
–
–
–
Xerocomellus
SOMF12854
MH011931
–
–
–
–
Xerocomellus
SOMF29860
MH011932
–
–
–
–
Xerocomellus
CM058
KP823760
–
–
–
–
Aureoboletus
pseudoauriporus
Farid 501
MW675741
MW662576
MW737500
MW737463
–
Aureoboletus
pseudoauriporus
JAB 124
MW675754
–
–
–
–
Aureoboletus
pseudoauriporus
JAB 130
MW675725
MW662581
–
–
–
Aureoboletus
pseudoauriporus
JAB 320
MW675726
MW662585
MW737508
MW737468
MW737489

38
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Aureoboletus
pseudoauriporus
JAB 80
MW675723
MW662588
MW737510
MW737471
MW737490
Cyanoboletus
bessettei
ARB 1393A
MW675734
MW662571
–
MW737457
MW737482
Cyanoboletus
bessettei
ARB 1393B
MW675735
–
–
MW737458
MW737483
Cyanoboletus
cyaneitinctus
JAB 324
MW675732
MW662586
MW737505
MW737469
–
Cyanoboletus
cyaneitinctus
JAB 325
MW675733
–
MW737506
MW737470
–
Cyanoboletus
cyaneitinctus
Farid 340
MW675739
MW662574
MW737502
MW737461
–
Cyanoboletus
cyaneitinctus
Farid 920
MW675744
MW662579
MW737503
MW737465
–
Cyanoboletus
cyaneitinctus
JAB 184
MW675731
MW662584
MW737504
MW737467
–
Cyanoboletus
cyaneitinctus
f. reticulatus
Farid 1035
MZ746113
–
–
–
–
Exsudoporus
floridanus
Farid 499
–
–
MW737497
MW737459
MW737484
Hemileccinum
floridanum
AB16
MW675745
MW662570
–
–
MW737481
Hemileccinum
floridanum
Farid 1032
MW675746
MW662573
–
–
–
Hemileccinum
floridanum
Farid 625
MW675742
MW662577
–
–
–
Hemileccinum
floridanum
JAB 142
MW675730
MW662583
–
–
MW737488
Lanmaoa
sublurida
Farid 1023
MW675736
MW662572
MW737498
MW737460
MW737485
Lanmaoa
sublurida
Farid 343
MW675740
MW662575
MW737499
MW737462
MW737486
Lanmaoa
sublurida
Farid 631
MW675743
MW662578
MW737501
MW737464
MW737487
Pulchroboletus
rubricitrinus
Farid 335
MF193884
MG026638
MW737512
MW737466
–
Xerocomellus
bolinii
JAB 110
MW675728
MW662580
MW737507
–
–
Xerocomellus
bolinii
JAB 43
MW675734
MW662587
MW737509
–
–
Xerocomellus
bolinii
JAB 133
MW675729
MW662582
–
–
–
Xerocomellus
bolinii
JAB 95
MW675735
MW662589
MW737511
MW737472
MW737491
Xerocomellus
salicicola
B391
MK552408
MW662569
MW737496
–
–
Hortiboletus
coccyginus
JLF 3093
KU144805
–
MW737513
MW737473
–
Xerocomellus
amylosporus
JLF 3498
KU144743
–
MW737514
MW737474
MW737492
Xerocomellus
rainisiae
JLF 3523
KU144789
KU144790
MW737515
MW737475
–
Xerocomellus
JLF 3558
KU144785
KU144786
MW737516
MW737476
–

39
Supplementary Table 1 Continued.
Species
GenBank
voucher
Locus
ITS
28S
RPB1
RPB2
TEF
Xerocomellus
atropurpureus
JLF 3620
KU144749
KU144750
MW737517
MW737477
MW737495
Xerocomellus
dryophilus
JLF 4134
KX534076
KY659593
–
MW737478
MW737493
Xerocomellus
dryophilus
JLF 4791
–
–
–
MW737479
MW737494
Xerocomellus
mendocinensis
JLF 5684
MH168533
MN294419
MW737518
MW737480
–
Xerocomellus
diffractus
JLF 5745
MH168534
–
MW737519
–
–