VOLUME 8

Fungal Systematics and Evolution DECEMBER 2021

PAGES 1–8

doi.org/10.3114/fuse.2021.08.01

Lentinula madagasikarensis sp. nov a elative o s ta e mus ooms om adagasca

B.P. Looney1, B. Buyck2, N. Menolli Jr.3,4, E. Randrianjohany5 . ibbett1*

1
Clark niversity e artment o Biology Worcester A 01 10 SA
2
us um national d istoire naturelle CN S Sor- bonne niversit Institut de Syst mati ue cologie Biodiversit IS EB E E rue Cuvier C
F- 00 aris France
3
Instituto Federal de Educa o Ci ncia e ecnologia de S o aulo IFS e artamento de Ci ncias da Nature a e atem tica C Sub rea de
Biologia SAB C m us S o aulo ua edro icente 2 S o aulo S 0110 -010 Bra il
4
Instituto de Bot nica IBt N cleo de es uisa em icologia N Av. iguel Ste ano gua Funda S o aulo S 04 01-012 Bra il
5
Centre National de echerche sur l Environnement CN E B 1 Lab. de icrobiologie de l Environnement L E Antananarivo adagascar

*Corresponding author: dhibbett clarku.edu

ey o ds st act We describe the rst s ecies o Lentinula from Africa, Lentinula madagasikarensis sp. nov. he ne ta on
Africa which was collected from central Madagascar, is strikingly similar to L. edodes, the shiitake mushroom. A BLAST search
biogeography using I S se uences rom L. madagasikarensis as the uery retrieves a mi o Lentinula, Gymnopus, Marasmiellus, and
edible mushrooms other members of Omphalotaceae as the top hits. A 28S phylogeny of the Omphalotaceae con rms lacement o L.
Omphalotaceae madagasikarensis within Lentinula. An I S hylogeny laces L. madagasikarensis as the sister group of L. aciculospora,
systematics which is a neotropical species. Lentinula madagasikarensis is characteri ed by robust basidiomata ith vinaceous ilei
1 ne ta on rominent occose scales near the ileus margin orets o s haero edunculate cheilocystidia and subcylindrical
basidios ores. his re ort constitutes a 4 000-mile trans-oceanic range e tension or Lentinula.

tation Looney B Buyck B enolli r N andrianjohany E ibbett 2021 . Lentinula madagasikarensis sp. nov. a relative o shiitake mushrooms
from Madagascar. Fungal Systematics and Evolution 8: 1–8. doi: 10.3114/fuse.2021.08.01
ece ved 14 April 2021; cce ted 3 May 2021; E ect vely u l s ed onl ne 1 June 2021
o es ond ng ed to P.W. Crous

INTRODUCTION analyses o I S se uences suggest that L. lateritia and L. edodes

Lentinula is a group of lignicolous agarics that includes the shiitake
mushroom, L. edodes. Shiitake is a traditional ood in East Asia
and it has re ortedly been cultivated in China or ca. 1 000 years
Chang iles 1 . o ever the genus Lentinula has a broad
distribution that s ans much o South Asia Australasia and tro ical
and subtropical regions of the Americas (Pegler 1983). Lentinula has
been resolved as a mono hyletic grou ithin the Omphalotaceae
(Agaricales), which also contains Gymnopus, Rhodocollybia, and
other collybioid mushrooms (Wilson & Desjardin 2005, Matheny
et al. 2006, He et al. 201 liviera et al. 2019).
Seven s ecies have been ormally described in Lentinula.
egler 1 included ve s ecies in his monogra h o Lentinula:
L. edodes in eastern Asia; L. lateritia in southeast Asia and
Australasia; L. novae-zelandiae in New Zealand; L. boryana
from the Gulf Coast of North America to South America; and L.
guarapiensis rom araguay. he latter is kno n only rom the ty e
collection S ega ini 1 . sing a combination o mor hology
mating com atibility and I S se uences ata etersen 2000
described a new species from Costa Rica, L. aciculospora. Mata
et al. 2001 then segregated a revious synonym Armillaria
raphanica, as L. raphanica from Pegler’s broad concept of L.
boryana. Lentinula raphanica is reported by Mata et al. (2001)
rom Florida Louisiana uerto ico Costa ica ene uela and
Bra il i.e. overla ing ith L. boryana s. str. . hylogenetic
may each contain multi le s ecies-level lineages but these have
not been ormally described ibbett et al. 1998).
Most species of Lentinula grow on wood of Fagales,
s eci cally Fagaceae or Nothofagaceae (Pegler 1983). The
absence of Lentinula from Europe and most of North America
here oaks and their relatives abound and outdoor log
cultivation o shiitake is success ul is u ling. istorical
biogeogra hic analyses have suggested that the resent
distribution o Lentinula could re ect an ancient trans-Beringian
distribution hich ould re uire local e tinction in North
America ibbett 2001 but alternative dis ersal routes via the
southern hemisphere cannot be rejected.
During a study of the wild edible mushrooms of Madagascar,
Buyck 200 made t o collections o a ungus that closely
resembled L. edodes. Here, we present morphological and
molecular analyses of the Malagasy material, which we describe
as L. madagasikarensis.
MATERIALS AND METHODS
Sam l ng and ylogenetic n e ence
o collections ere made by Buyck 200 in native orests
of central Madagascar with endemic Uapaca densifolia

Fungal Systematics and Evolution is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License
© 2021 Westerdijk Fungal Biodiversity Institute 1
 Looney et al.
(Phyllanthaceae) and Sarcolaenaceae, and introduced
Eucalyptus robusta in 200 and 200 . Both collections have
been de osited at the ungarium C o the us um National
d istoire Naturelle aris. enomic NA as e tracted rom
both collections in 2020 using the E traction Solution-based
method of Looney et al. (2020).
o nuclear loci ere am li ed and se uenced ith standard
rimer sets I S1F-I S4 White et al. 1 0 or I S and L -L
ilgalys ester 1 0 or 2 S. Both strands ere se uenced
using the Sanger method by somagen Ne ork N and
se uences have been de osited into enBank accession Nos.
MW810299–MW810302).
o datasets ere used to in er relationshi s at di erent
scales, Dataset 1 was used to assess the placement within the
Omphalotaceae and Dataset 2 was used to infer placement
within Lentinula. Dataset 1 used the 28S Omphalotaceae matri
rom liveira et al. 201 as a starting alignment. ataset 2
included 0 I S se uences re resenting all kno n lineages o
Lentinula, which were downloaded using emerencia (Nilsson et
al. 200 and si se uences o Gymnopus spp. as the outgroup.
Se uences o the adagascar material ere manually
aligned ith the t o datasets in Ali ie v. 1.2 Larsson
2014 . hylogenetic in erence under a imum Likelihood as
er ormed in A L- I- C using ra ml I v. 2.0 ith auto E
bootstopping criteria (Stamatakis 2006, Edler et al. 2020). Bayesian
analysis as er ormed in rBayes v. .2. a using etro olis
cou led C C under a model through the CI ES
Science gate ay on uist et al. 2012). The analysis consisted
o runs o 0 generations and our chains sam led every
000 generations. Finally a distance matri o nine re resentative
se uences o di erent ta a as generated rom the nal I S
alignment in using the dist.dna unction o the a e ackage.
Alignments and resulting trees have been de osited in reeBASE
(htt s treebase.org ) under study number 28094.
o olog cal analys s
or hological eld notes ere taken on resh basidiomata using
color standards given in arentheses orneru Wanscher
1 . icrosco ic observations ere made on a Nikon
Eclipse e600 compound microscope with a mounted SPOT RT
Slider digital camera. Dried specimens were rehydrated in 3 %
and observed in hase contrast or stained ith hlo ine
and by bright eld microsco y. el er s reagent as used or
testing amyloidity. At least 20 observations ere made or each
eature. icrosco ic eature measurements are given as lo er
limit arithmetic mean u er limit ith n = total number
o observations and -value or basidios ores calculated as
arithmetic mean o the lengths divided by arithmetic mean o
the idths or the lo er limit arithmetic mean u er limit.
RESULTS
olecula systematics
BLAS searches using I S se uences rom L. madagasikarensis
as the ueries retrieved se uences o Lentinula, Gymnopus,
Marasmiellus, and other members of Omphalotaceae as the
to hits results not sho n . hylogenetic analyses o ataset
1 recovered all o the major clades o Omphalotaceae identi ed
by liveira et al. 201 but o en ith eak bootstra su ort
2 © 2021 Westerdijk Fungal Biodiversity Institute
(BS) (Fig. 1). Lentinula as strongly su orted as mono hyletic
(BS 91 %). The sister group of Lentinula as not resolved ith
con dence but a clade containing Lentinula and Clades A–J of
liveira et al. (2019) was strongly supported (BS 99 %).
The Lentinula- ocused I S analyses o ataset 2 laced L.
madagasikarensis as sister to L. aciculospora with moderate BS
4 Fig. 2 . o major grou s ere resolved in Lentinula,
one eakly-su orted grou containing all Asian-Australasian
s ecies BS and one strongly-su orted clade containing
all American species, plus L. madagasikarensis (BS 98 %). There
are three major groups in the Americas, L. aciculospora (BS
100 %), L. boryana (BS 86 %), and L. raphanica (BS 100 %).
he Asian-Australasian clade contains ve lineages that have
reviously been termed rou s 1 based on I S data ibbett
et al. 1 . Each o the I S-based grou s are moderately to
strongly su orted as mono hyletic but relationshi s among
them are not ell resolved Fig. 2 . rou s 1 and corres ond
to L. edodes sensu lato including cultivated shiitake mushrooms.
Based on sam ling in the resent study and that o ibbett et
al. (1998), Group 1 includes isolates from China, Japan, South
orea hili ines India and the ussian Far East hereas
rou includes isolates rom China Ne al and India. rou 2
and rou 4 both include collections rom a ua Ne uinea
rou 2 also includes collections rom Australia. rou is
e uivalent to L. novae-zelandieae, from New Zealand.
air ise distances o I S se uences o L. madagasikarensis to
other species of Lentinula ranged from 11 % vs. L. edodes (Group
5), L. lateritia (Group 2), L. novae-zelandiae, and L. aciculospora,
to 20 % vs. L. raphanica Fig. . air ise se uence distances
among three isolates from the Americas are 11–20 %, whereas
those among ve Asian-Australasian isolates are 1 4 Fig. .
Taxonomy
Lentinula madagasikarensis Buyck, Randrianjohany & Looney,
sp. nov. MycoBank MB 839129. Fig. 4A–G.
Etymology he s eci c e ithet is derived rom adagasikara
which is the Malagasy name for Madagascar. To our knowledge,
there is no Malagasy name for this species.
Diagnosis obust basidiomata ith vinaceous ileus
color and large tu ed scales near ileus margin merging
into an a endiculate margin. Fibrous ivory-colored
sti e ith s uamules. Small and narro basidios ores.
S haero edunculate cheilocystidia orming scattered orets.
Typus: adagasca oramanga district Alaotra- angoro
region, Andasibe, 18°56’S 48°25’E, on E. robusta log, 20 Jan.
2006, B. Buyck & V. Hofstetter 06.007 (holotype PC0142531).
Description: Pileus conve to hemis haerical hen young soon
umbonate to a lanate or broadly de ressed rm and eshy
up to 6– cm diam. sur ace smooth sometimes scur y or
with minute scales or granules and almost greasy at the touch
in the central ortion to ard the margin ra idly becoming
covered ith a aler hairy- brillose scur building more or less
concentrically arranged thick occose de osits ith brillose
e tremities ointing do n ard dark reddish bro n EF - to
vinaceous or ur lish bro n 14EF - at center hen young
at maturity becoming paler, pale reddish brown to ochraceous
bro n A - 4B - C - but retaining locally some darker
 Lentinula madagasikarensis sp. nov.

Clade A 83 seqs
Marasmiellus s.str.
Lentinula raphanica INPA1701G
99
87 Lentinula raphanica TFB8682-2
81 Lentinula boryana TFB10292-6
99 Lentinula madagasikarensis sp. nov BB06007 Clade F
91 Lentinula madagasikarensis sp. nov BB08120
100
1 0 0 Lentinula edodes OE9
Lentinula edodes Le Bin 0899
Lentinula
Lentinula lateritia DSH 9214

100 Clade B 18 seqs

Pusillomyces

Clade D 13 seqs
Pallidocephalus

Clade E 12 seqs
Rhodocollybia
52
100 Clade C 6 seqs
Connopus

54 69 Clade H 54 seqs
73 63
Paragymnopus
99
86 Clade G 51 seqs
Gymnopus s. str.
KY026621 Marasmius sp. 1 TFB3940

66
97 Clade I 7 seqs
Gymnopanella

Clade J 10 seqs
Mycetinus

100
Clade K Omphalotus 2 seqs
98
92
91
99
89 Outgroups
66 100

0.03

Fig. 1. a imum-likelihood hylogeny o Omphalotaceae based on the 2 S se uence dataset o liveira et al. (2019) with 788 bootstrap replicates
according to autoMRE bootstopping criteria. Major clades from the original study are labeled and collapsed with number of samples included in each
clade. BS values 0 are included ith the branches.

shades. Lamellae adnato- or adne o-sinuate ith a decurrent
tooth later re uently sub ree une ual ith u to 4–5 series
o lamellulae o di erent lengths hitish to ivory discolouring
bro nish a er injury cro ded edge concolorous entire or
irregularly serrulate. Stipe up to 4 × 2.5 cm; stout and rm al ays
shorter than the pileus diameter, central to mostly eccentric,
cylindrical or a little idening to almost bulbous near the base
strongly brillose-s uamose over the entire sur ace rom thick
brils or s uamae ointing out- or u ard solid. Partial veil a
thick cottony tissue strongly brillose-s uamose on the lo er
sur ace rom brils ointing do n ard ugacious soon breaking
u and limited to remnants on the ileal margin or sometimes
leaving a artial ring on u er art o the sti e. Context rm-
eshy ale coloured not discolouring but o en colored yello ish
in the sti e near the attachment to the oody host sur ace.
Spore print white. Basidiospores oblong to subcylindrical, 5.0–
.0 .0 2.0 .0 . m 1. 2.1 2.4 n = 30), hyaline,
inamyloid thin- alled smooth hilar a endage cons icuous
contents heterogenous or homogeneous. Hymenium composed
mostly of basidia and basidioles. Basidia 11.0 1 .0 2

© 2021 Westerdijk Fungal Biodiversity Institute 3

 Looney et al.

U33082 L. novaezelandiae G3 TMI1449 NZL

U33079 L. novaezelandiae G3 RHP7563 NZL
93/1 U33075 L. novaezelandiae G3 NZFS210 NZL
U33074 L. novaezelandiae G3 NZFS156 NZL
L. novaezelandiae Group 3
AB366151 L. novaezelandiae G3 as L. edodes D703PP9 NZL New Zealand
U33081 L. novaezelandiae G3 TMI1172 NZL
U33073 L. lateritia G2 DSH92149 PNG
AF079574 L. lateritia G2 wc802 PNG
AF031180 L. lateritia G2 RV95377 AUS
60/0.99 U33071 L. lateritia G2 DSH92145 PNG
U33076 L. lateritia G2 RHP3577 AUS
1 U33072 L. lateritia G2 DSH92147 PNG
79/0.99 U33083 L. lateritia G2 TMI1476 PNG
L. lateritia Group 2
Asian - AF031179 L. lateritia G2 RV95376 AUS Australia & Papua New Guinea
AF031181 L. lateritia G2 RV95378 AUS
Australasian AF031192 L. lateritia G2 DSH92-148 PNG
AF079573 L. lateritia G2 wc800 PNG
clade U33084 L. lateritia G2 TMI1485 PNG
54/0.88 AF031191 L. edodes G5 TMI1546 NPL
94/1 LC149603 L. edodes G5 B2SN033 NPL L. edodes Group 5
LC149606 L. edodes G5 B2SN037 NPL Asia
56/0.97 AF356169 L. edodes G5 as L. lateritia BDhanukasn IND
100/1 U33086 L. lateritia G4 TMI1502 PNG L. lateritia Group 4
U33085 L. lateritia G4 TMI1499_PNG
AY636053 L. edodes G1 OE9 PHL Papua New Guinea
MG735346 L. edodes G1 LeBIN0899 RUS
JX984748 L. edodes G1 as Uncultured fungus FA1021 KOR
EF174453 L. edodes G1 UASWS0314 POL
99/1 KM985682 L. edodes G1 as L. sp. BPSM45 IND L. edodes Group 1
AY636052 L. edodes G1 OE2 IND Asia
AF356170 L. edodes G1 LeBIN1024 RUS
EF174451 L. edodes G1 UASWS0311 POL
AF079579 L. raphanica as L. boryana sp834 BRA
100/1 AF079578 L. raphanica as L. boryana LB4264 VEN
100/1 AF356168 L. raphanica DSH98003 PRI
AF031178 L. raphanica as L. boryana HN2002 USA
AY016441 L. raphanica TFB9156 USA L. raphanica
AY256687 L. raphanica TENN56663 CRI Americas
99/1 KF937345 L. raphanica AMV1850 COL
KY026653 L. raphanica TFB8682 USA
AY016442 L. raphanica TFB9564 PRI
86/1 AY016440 L. boryana TFB10827 BRA
KY026657 L. boryana TFB10292 MEX
AF031175 L. boryana RGT96062409 CRI
97/0.99 AF031175 L. boryana RGT960624 CRI
98/1
U33078 L. boryana R52 MEX L. boryana
U33077 L. boryana R39 MEX Americas
AF031177 L. boryana IE154R50 MEX
American - AF031176 L. boryana IE17R38 MEX
African 100/1
AF079576 L. boryana wc794 MEX
L. madagasikarensis sp. nov.
L. madagasikarensis BB06.007 MAD HOLOTYPE
clade L. madagasikarensis BB08.120 MAD Madagascar
100/1 JQ247977 L. aciculospora PPNag001 EQU
84/0.99
AY016444 L. aciculospora TFB10418 CRI
L. aciculospora
AY016443 L. aciculospora TFB9447 CRI HOLOTYPE Central America
JX536172 G. aquosus BRNM665362
JX536141 G. dryophilus BRNM670778
DQ241781 G. dryophilus AFTOL559
100/1
JX536139 G. dryophilus BRNM737691 Outgroup
JX536157 G. dryophilus BRNM707149
JX536146 G. alpinus CB16251

0.05

Fig. 2. a imum-likelihood hylogeny o Lentinula based on ataset 2 o I S. BS values 0 Bayesian values 0. 0 are included ith the
branches. Se uence labels include enBank accession numbers and collection numbers. 1 in the Asian-Australasian clade corres ond to I S-
based grou s rom ibbett et al. 1 . Country codes A S Australia B A Bra il C L Colombia C I Costa ica IN India South
orea E e ico N L Ne al N L Ne ealand L hili ines N a ua Ne uinea L oland ossible cultivar I uerto
ico S ussia SA continental SA EN ene uela.

4.0 .0 . m n 40 oblong to cylindrical clavate ith a
median constriction ith a clam connection at the base our-
sterigmate sterigmata slender u to m long. Pleurocystidia
absent. Cheilocystidia 1 .0 2 4 .0 11.0 1 .0 m n =
20 clavate to s haero edunculate in ated a ically ithout
lobes thin- alled smooth orming dense clusters or orets
in re uent. Lamellar trama regular to subregular; hyphae 4.0–
1 .0 m diam. thin- to thick- alled. Subhymenium rudimentary.
Epicutis 30–50 m thick hy hae .0 .0 m diam bro n in
mass hyaline singly re ent inter oven and subregular cutis
hyphae in scales erumpent, ending in eroded hyphal fragments.
Subpellis com osed o re uently branching irregular hy hae
.0 11.0 m diam hyaline thick- alled. Stipitipellis composed
o inter oven hy hae .0 .0 m diam branching ith occons
formed of obtuse terminal end cells. Caulocystidia absent.

4 © 2021 Westerdijk Fungal Biodiversity Institute

 Lentinula madagasikarensis sp. nov.

L. raphanica 0.20 0.20 0.21 0.20 0.21 0.20 0.20 0.17 0.00

L. boryana 0.12 0.12 0.12 0.12 0.12 0.14 0.15 0.00 0.17

L. aciculospora 0.12 0.11 0.12 0.10 0.11 0.11 0.00 0.15 0.20

L. madagasikarensis 0.12 0.11 0.12 0.11 0.11 0.00 0.11 0.14 0.20

L. lateritia G2 0.03 0.01 0.03 0.01 0.00 0.11 0.11 0.12 0.21

L. edodes G5 0.02 0.01 0.03 0.00 0.01 0.11 0.10 0.12 0.20

L. lateritia G4 0.04 0.03 0.00 0.03 0.03 0.12 0.12 0.12 0.21

L. novaezelandiae 0.03 0.00 0.03 0.01 0.01 0.11 0.11 0.12 0.20

L. edodes G1 0.00 0.03 0.04 0.02 0.03 0.12 0.12 0.12 0.20
s
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ia

ia
es
an

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Fig. 3. istance-matri heat ma o re resentative se uences o s ecies o Lentinula based on air ise distances rom the I S multi-se uence
alignment given as ro ortion o divergence.

Habit and habitat regarious to scattered on corticate logs o E.
robusta and unidenti ed hard ood in humid mi ed mountain
orests at higher elevations o the central lateau and eastern
escarpment of Madagascar.
Additional specimen examined: adagasca Anka obe district
Analamanga region Ambohitantely S ecial eserve 1 .1 1 S 4 . 02 E
in primary forest dominated by U. densifolia and Sarcolaenaceae, 22
Jan. 2008, B. Buyck & V. Hofstetter no. 08.120 (paratype PC0142532).
Notes: Lentinula madagasikarensis can readily be distinguished
from its apparent sister species, L. aciculospora, by gross
mor hological characters including a darker vinaceous
pileus color in L. madagasikarensis and larger tu ed scales
concentrated near the ileus margin orming a distinctive
a endiculate margin as ell as microsco ic di erences in
basidios ore si e hich is smaller in L. aciculospora (5.6–8.8
1. 2. m and sha e o cheilocystidia that are gnarled
and bluntly lobed in L. aciculospora (Mata & Petersen 2000).
Lentinula madagasikarensis closely resembles L. boryana (Mata
et al. 2001 but di ers microsco ically because the rst never
has a endages on cheilocystidia and caulocystidia are absent.
Macroscopically, L. madagasikarensis di ers rom L. boryana
in si e color and robustness o the ilei hich are smaller
(1.8–2.5 cm) and paler (light brown to golden brown or grayish
orange) in L. boryana (Mata et al. 2001). Morphologically, L.
madagasikarensis closely resembles L. edodes in pileus color
and the robust aspect of the basidiomata, which also has a
dark vinaceous bro n and cm diam ileus egler 1 .
Microscopic characters are similar between the two species
ith similar si e o the basidios ores . . m in L.
edodes and general sha e o cheilocystidia also clavate ithout
any apical appendages in L. edodes egler 1 . o ever
basidiospores of L. edodes are slightly more ovoid 1. as
re orted by egler 1 and cheilocystidia are not re orted as
s haero edunculate or orming orets as is ty ically ound in
L. madagasikarensis. he geogra hic distribution also se arates
L. edodes from L. madagasikarensis ith the rst broadly
distributed in Asia and the latter kno n only rom adagascar.

© 2021 Westerdijk Fungal Biodiversity Institute 5

 Looney et al.

Fig. 4. A, B. Field photograph of Lentinula madagasikarensis (type specimen, PC coll. BB06.007). C, D. Field photographs of Lentinula madagasikarensis
(paratype specimen, PC coll. BB08.120). E. Basidiospores. F. Basidia. G. Cheilocystidia. Scale bars A 2 cm E 20 m.

6 © 2021 Westerdijk Fungal Biodiversity Institute

 Lentinula madagasikarensis sp. nov.
DISCUSSION
Buyck (2008: 516) described the Malagasy Lentinula as a
shiitake look-alike. ur analyses con rm that his collections
rom t o di erent localities in adagascar re resent a
new species of Lentinula. Lentinula madagasikarensis was
discovered 4 000 miles rom the nearest ell-documented ild
o ulations o Lentinula, which are those of L. edodes Group
in India and Ne al. Its a arent sister grou L. aciculospora,
occurs 9 000 miles away, in Costa Rica (Mata & Petersen 2000),
Ecuador (Andrade et al. 2012), Panama (Piepenbring 2008), and
Nicaragua me 201 . o ever several collections identi ed
as L. edodes or “L. edodes-like have been re orted rom the
emocratic e ublic o the Congo C hich is less than 2 000
miles rom adagascar lobal Biodiversity In ormation Facility
.gbi .org occurrence records 1 2 4 1 40 410
1 40 444 . As o this riting e have not been able to study
the material from the DRC.
At its narro est oint the o ambi ue Channel se arates
Madagascar and mainland Africa by less than 300 miles. Based
on their geogra hic ro imity one ould e ect a Lentinula
species from the DRC and L. madagasikarensis to be closely
related. Even i they are the resence o Lentinula in Madagascar
is robably due to long-distance dis ersal adagascar began to
se arate rom continental A rica during the Late urassic eriod
about 1 0- -yr ago de Wit 200 hereas Lentinula is about
0- -yr-old according to a molecular clock analysis by arga et
al. (2019).
Both collections o L. madagasikarensis were found in similar
habitats in the highlands of the Central Plateau at 1 500–2 000 m
alt. ne collection as ound gro ing on ood o E. robusta and
the other as on a allen log o an unidenti ed native tree Buyck
2008: 516), but both were in dense forests largely dominated by
U. densifolia and various s ecies o Sarcolaenaceae. All other
s ecies or s ecies com le es o Lentinula are reported to occur
on Fagales articularly Fagaceae or Nothofagaceae, but some
are also capable of growing on other hardwood substrates
egler 1 ata etersen 2000 c en ie et al. 2000, Mata
et al. 2001 ie enbring 200 . here are no native Fagaceae or
Nothofagaceae in adagascar or Sub-Saharan A rica anos
Stan ord 2001 na et al. 200 . A recent discovery o 2- -yr-
old Castanopsis fossils from Patagonia shows that Fagaceae was
present in the southern hemisphere in the early Eocene (Wilf et
al. 2019), but that was late in the breakup of Gondwana, long
a er the o ening o the South Atlantic cean. Nothofagus has
an 0- -yr ossil record but there is no evidence o this grou
having e isted in A rica or adagascar na et al. 2005).
he hylogenetic lacement o L. madagasikarensis
is im ortant or understanding host shi s and historical
biogeography of Lentinula. he distribution o host ranges in
Lentinula suggests that the ancestor of the genus decayed
Fagales. I so then L. madagasikarensis would represent an
e ansion onto hosts that are indigenous to adagascar as
ell as introduced eucaly ts. Based on the I S hylogeny L.
madagasikarensis is closely related to L. aciculospora within the
American-A rican clade Fig. 2 suggesting that it and ossibly
the entire genus is derived rom an ancestor in the neotro ics.
o ever many branches in the I S hylogeny o Lentinula
are not strongly resolved Fig. 2 . We de er ormal analyses o
historical biogeogra hy and evolution o substrate ranges until
e have a genome se uence or L. madagasikarensis, which we
are currently pursuing.
© 2021 Westerdijk Fungal Biodiversity Institute
ACKNOWLEDGEMENTS
BB e resses sincere thanks to the Committee or esearch and
E loration o the National eogra hic Society or unding art o
the eld ork through grant 21-0 . Colleagues o the CNRE at
Antananarivo are ackno ledged or e cellent eld assistance. adson .
S. liveira kindly rovided the starting alignment or ataset 1. esearch
at Clark niversity as su orted by National Science Foundation
a ard EB-14 S I.
on ct o nte est he authors declare that there is no con ict
of interest.
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