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Sequestrate species of Agaricus and Macrolepiota from Australia: New species and combinations and their position in a calibrated phylogeny

Mycologia

November 2011
104(2):496-520

DOI:10.3852/11-092

Source
PubMed

Authors:

Teresa Lebel

Anna E Syme

University of Melbourne

Australian collections of sequestrate Agaricaceae were examined with morphological and molecular data (nuclear DNA from ITS and LSU), and the majority were found to belong to the genera Agaricus and Macrolepiota. Previously described Australian species of Endoptychum are transferred to the appropriate agaricoid genera and several new combinations proposed. Descriptions and illustrations are provided for these and eight new species: Agaricus eburneocanus sp. nov., A. chartaceus sp. nov., A. erythrosarx sp. nov., A. inilleasper sp. nov., A. pachydermus sp. nov., Macrolepiota gasteroidea sp. nov., M. vinaceofibrillosa sp. nov. and M. turbinata sp. nov. The sequestrate genus Barcheria is retained as a distinct taxon. Timing of evolution of sequestrate sporocarp forms in Macrolepiota, Chlorophyllum and Agaricus seems to have occurred in the past 15 000 000 y, and a stem age is approximately 65 000 000 y for Barcheria.

Bayesian phylogram of Chlorophyllum and Macrolepiota based on ITS data. Thick branches indicate support by posterior probabilities of. 0.95. All sequestrate taxa are in boldface.

…

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Content may be subject to copyright.

Sequestrate species of

Agaricus

and

Macrolepiota

from Australia:

new species and combinations and their position in a calibrated phylogeny

Teresa Lebel

Anna Syme

National Herbarium of Victoria, Royal Botanic Gardens

Melbourne, Private Bag 2000, Victoria, 3141 Australia

Abstract

: Australian collections of sequestrate Agar-

icaceae were examined with morphological and

molecular data (nuclear DNA from ITS and LSU),

and the majority were found to belong to the genera

Agaricus

and

Macrolepiota

. Previously described Aus-

tralian species of

Endoptychum

are transferred to the

appropriate agaricoid genera and several new combi-

nations proposed. Descriptions and illustrations are

provided for these and eight new species:

Agaricus

eburneocanus

sp. nov.,

A. chartaceus

sp. nov.,

erythrosarx

sp. nov.,

A. inilleasper

sp. nov.,

pachydermus

sp. nov.,

Macrolepiota gasteroidea

sp.

nov.,

M. vinaceofibrillosa

sp. nov. and

M. turbinata

sp. nov. The sequestrate genus

Barcheria

is retained as

a distinct taxon. Timing of evolution of sequestrate

sporocarp forms in

Macrolepiota

Chlorophyllum

and

Agaricus

seems to have occurred in the past

15 000 000 y, and a stem age is approximately

65 000 000 y for

Barcheria

Key words:

Agaricaceae,

Endoptychum

, nomencla-

ture, taxonomy, timing of evolution

INTRODUCTION

The Agaricaceae as currently understood includes

taxa with considerable diversity in sporocarp form,

from agaricoid to sequestrate (hypogeal or emergent,

hymenophore remaining enclosed, and lacking active

spore dispersal, and now pulverant gastroid (Singer

1986, Candusso and Lanzoni 1990, Redhead et al.

2001, Kruger et al. 2001, Kirk et al. 2008). Studies

have shown that sequestrate sporocarp forms have

evolved relatively recently, with up to 10 independant

gastromycetization or sequestration events scattered

throughout the family (Kru¨ger et al. 2001, Moncalvo

et al. 2002, Vellinga 2004, Lebel et al. 2004, Matheny

et al. 2007, Gube 2009). Historically sequestrate taxa

were placed in morphologically distinct genera, such

Endoptychum

(Czernajew 1845),

Gyrophragmium

(Montagne 1843),

Montagnea

(Fries 1836) or

Barcheria

(Lebel et al. 2004) and until fairly recently were treated

in unique families (e.g. Julich 1982:

Podaxaceae

Montagnaceae

Secotiaceae

The genera

Agaricus

Chlorophyllum

and

Macro-

lepiota

are easily recognized based on sporocarp

morphology and a few microscopic characters, how-

ever species differentiation can be problematic

(Vellinga 2003, Geml et al. 2004, Kerrigan et al.

2006, Ge et al. 2010). All three genera are important

saprotrophs in many habitats from the tropics to the

arctic, and several edible and toxic species can be

found in each genus (Ding and Huang 2003, Kirk

et al. 2008).

Agaricus

is the most species rich with more

than 300 known worldwide, then

Macrolepiota

with 30

species and

Chlorophyllum

with 16 species (Bas 1991,

Kirk et al. 2008). The Australian taxa in these lineages

are generally understudied and poorly known. Of the

37 currently known species of

Agaricus

, more than half

are introduced taxa (Grgurinovic 1997, May et al.

2006). Neither

Macrolepiota

nor

Chlorophyllum

are

particularly diverse in Australia, with three and four

species respectively (Vellinga 2003).

Of the sequestrate taxa with affinities to

Agaricus

the North American species

Longula texensis

(Berk.

& M.A. Curtis) Zeller (5

A. deserticola

G. Moreno,

Esqueda & Liza´rraga),

Gyrophragmium dunalii

Zeller

A. aridicola

Geml, Geiser & Royse) and

Endopty-

chum depressum

Singer & A.H. Sm.(5

A. inapertus

Vellinga) are members of

Agaricus

section

Arvenses

(Moncalvo et al. 2000; Geml 2004, 2004; Gube 2009).

The relationships of Australian taxa

Secotium coarcta-

tum

Berk.,

Endoptychum wariatodes

Grgur.,

E. moon-

gum

Grgur. and

E. melanosporum

(Berk.) Singer &

A.H. Sm. also are likely to be with

Agaricus

, based on

morphology. The rare taxon

Smithiogaster volvoagar-

icus

J.E. Wright appears to be an aborted

Agaricus

(Wright 1975). Finally

Endoptychum agaricoides

Czern. was found to belong to

Chlorophyllum

agaricoides

[Czern.] Vellinga), but the position of

arizonicum

is unresolved, being sister to

Agaricus

and

Chlorophyllum

(Vellinga 2002, Gube 2009). No se-

questrate taxa related to

Macrolepiot a

have been

described thus far.

Based on extensive morphological examination

and molecular phylogenetic analyses, we present

species diversity and descriptions of novel sequestrate

Agaricus

and

Macrolepiota

from Australia and their

affinities to other species in these lineages. In

addition we conducted a trial calibrated phylogenetic

Submitted 23 Mar 2011; accepted for publication 13 Aug 2011.

Corresponding author. E-mail: teresa.lebel@rbg.vic.gov.au

E-mail: anna.syme@rbg.vic.gov.au

Mycologia,

104(2), 2012, pp. 496–520. DOI: 10.3852/11-092

2012 by The Mycological Society of America, Lawrence, KS 66044-8897

496

analysis, using existing hypotheses of estimated taxon

divergences to provide preliminary evidence of the

timing of evolution of sequestrate sporocarp forms in

the three lineages of

Agaricus

Chlorophyllum

and

Macrolepiota

(Geml et al. 2004).

MATERIALS AND METHODS

Taxon sampling.—

The regions of nuclear rDNA data used

for analyses included the internal transcribed spacers ITS1–

5.8S–ITS2 (ITS) and the 59 end of the large subunit (LSU).

Sequences representing 127 species within Agaricaceae

were derived from fresh or herbarium material or were

retrieved from GenBank. Novel sequences representing

Australian species of

Agaricus

(40),

Barcheria

(seven) and

Macrolepiota

(11), and three of

Chlorophyllum agaricoides

were generated for this study (SUPPLEMENTARY TABLE I).

Unfortunatelywewereunabletoobtainmaterialor

sequences of

E. arizonicum

for our analyses. Type material

Endoptychum melanosporum

Secotium coarctatum

and

wariatodes

is fragmentary and has been stored in an

unknown chemical; no useable DNA was obtained.

Nucleic acid preparation, amplification and sequencing.—

Genomic DNA was isolated with the QIAGEN DNeasy Plant

Mini Kit, following the manufacturer’s protocol. The

targeted regions were amplified from purified DNA with

standard fungal primer pairs ITS1/ITS4 (White et al. 1990,

Gardes and Bruns 1993) or LROR/LR5 (http://www.

biology.duke.edu/fungi/mycolab/primers.htm).

PCR protocols followed standard methods (Lebel et al.

2004, Lebel and Tonkin 2007, Danks and Lebel 2010).

Purified DNA (Concert Rapid PCR Purification System, Life

Technologies) was directly sequenced with the ABI Prism

BigDye Terminator Cycle Sequencing Kit with primers ITS1

and ITS4 and LROR and LR5. Sequencing was carried out

with an ABI model Automated 377 DNA Sequencer.

Assembly and manual editing of sequences for each region

were performed with Sequencher 4.3 (Gene Codes Corp.).

Sequences were aligned with default settings in MUSCLE

(Edgar 2004) with some manual editing. Previously unpub-

lished sequences are deposited in GenBank (S

UPPLEMENTA-

RY TABLE

I).

Alignments and phylogenetic analyses.—

A preliminary align-

ment consisting of 72 LSU sequences and 965 characters

was generated to investigate broad patterns of relationships

between genera. Subsequently two alignments were gener-

ated to investigate positions and relationships within (a)

Agaricus

, consisting of 102 ITS sequences and 854

characters and (b)

Macrolepiota

and

Chlorophyllum

, consist-

ing of 87 ITS sequences and 896 characters. Gaps in

alignments were treated as missing data.

Tulostoma

squamosum

was included as outgroup in all alignments

based on work showing it as a potential sister taxa to these

lineages (Vellinga et al. 2003, Vellinga 2004, Matheny et al.

2007, Gube 2009). A combined ITS-LSU analysis was not

undertaken because LSU data was not available for the

majority of taxa. All alignments and trees are available

in TreeBase (http://purl.org/phylo/treebase/phylows/

study/TB2:S11712; temporary reviewer access http://purl.org/

phylo/treebase/phylows/study/TB2:S11712?x-access-code5

6280681c731f036e5967bd14672529b8&format5html).

Phylogenetic analyses were performed with Bayesian

likelihood with the Metropolis coupled Markov chain

Monte Carlo (MCMC) search algorithm implemented in

MrBayes 3.1.2 (Ronquist and Huelsenbeck 2003). PAUP*

4.0b10 (Swofford 2002) and Modeltest (Posada and

Crandall 1998) were used to determine appropriate

evolutionary models for the Bayesian analyses. Under the

Akaike information criterion (AIC) the best-fit models for

each alignment had six substitution types (nst 5 6), gamma-

shaped rate variation and a proportion of invariable sites

(rates 5 invgamma). Each search involved four chains

(three heated to 0.2 and one cold) sampled every 1000

generations. The search ran until convergence between

runs was achieved, measured by a standard deviation of split

frequencies below 0.05 and observing when this was no

longer decreasing. For ITS

Agaricus

this was 0.02 after

5 000 000 generations; for ITS

Chlorophyllum

Macrolepiota

0.01 after 5 000 000 generations. To confirm that the

topologies had converged we visualized split posteriors with

AWTY (Wilgenbusch et al. 2004). After discarding burn-in,

consensus trees were imported to FigTree (Rambaut 2009)

and are presented with posterior probabilities at clades.

Calibration of phylogeny.—

We used a combined subset of

the ITS alignments, with a focus on lineages with both

sequestrate and agaricoid sporocarp forms within the three

lineages of interest to decrease computational time (taxa

included are annotated by asterisk in S

UPPLEMENTARY TABLE

I). To construct the calibrated phylogeny we used the

program BEAST (Drummond and Rambaut 2007). An XML

input file was created with the ancillary program BEAUTi.

The substitution model was general time reversible (GTR)

with + I + G as found by Modeltest as described earlier. We

used the uncorrelated lognormal (UCLN) relaxed clock

model for nucleotide evolution, which permits among-

lineage rate heterogeneity to be uncorrelated and performs

well whether the data are clock-like or not (Drummond

et al. 2006). A random starting tree was used and a Yule

model for the tree prior, a model that is useful for species-

level analyses (Drummond and Rambaut 2007).

For the most recent common ancestor (MRCA) of

Agaricus

and

Chlorophyllum

we placed a prior distribution

on the divergence time of

A. xanthodermus

and

C. molybdites

of 73 000 000 y ago (Geml et al. 2004), based on a multigene

study of Heckman et al. (2001). Several distributions can be

chosen to model the prior, but a normal distribution is

useful for modelling the uncertainty around secondary

calibration points such as this one (Ho 2007). We therefore

used a normal prior on this divergence time and set the

value of the mean at 73 and a standard deviation of 1.0

(making 95% of the probability 71 000 000–75 000 000 y).

Two independent runs of 20 000 000 generations were set,

sampling every 2000 steps. The program Tracer was used to

check that the effective sample size (ESS) was greater than

or close to 200 for the posterior, prior and likelihood,

indicating that samples are independent (Drummond and

Rambaut 2007). A burn-in of 10% was found to be adequate

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

497

as stationarity had been achieved, and the distributions

across the separate runs were congruent. The two runs were

combined with LogCombiner, each with a 10% burn-in

(1000 trees) and the parameters and topology summarized

in TreeAnnotator (Drummond and Rambaut 2007). The

maximum credibility tree (MCC) with node heights set to

mean heights was imported to FigTree (Rambaut 2009) and

plotted against a time scale.

We also constructed a binary character matrix for

morphology of sequestrate forms as absent (0) or present

(1) (S

UPPLEMENTARY TABLE II). With Mesquite (Maddison

and Maddison 2010) this character was mapped onto the

calibrated phylogeny with the Markov k-state 1-parameter

model (MK1), which assumes equal probabilites for change

between states.

Morphology.—

Macroscopic characters were described di-

rectly from fresh material and based on examination of,

and field notes on, dried collections. Colors are described

in general terms. Habitat, associated plant communities

and fruiting season are based on collection notes. Hand-

cut sections of fresh and dried material were mounted

either in 5% aqueous solution of KOH then stained with

ammoniacal Congo red or cotton blue or directly stained

with Melzer’s reagent. Measurements were made at 4003

or 10003 with a calibrated ocular micrometer. Spore

dimensions are given as length range 3 width range (n 5

20). The length : width ratio (Q) of individual spores is

presented as the range of Q values. Measurements do not

include the apiculus. Dimensions of basidia and cystidia

are given as length range 3 width range (n 5 15). All

drawings and illustrations are based on type material unless

otherwise stated. Informal taxon names are standardized

according to Barker (2005) and names of herbaria are

abbreviated according to Thiers (http://sweetgum.nybg.

org/ih/ continuously updated).

RESULTS

Phylogenetic analyses.—Agaricus ITS dataset. Microp-

salliota

(including

Allopsalliota geesteranii

)and

Chloro-

phyllum

both form well supported clades (FIG.1).If

PPs of , 0.95 are collapsed,

Heinemannomyces

Clarkeinda

Eriocybe

and

Coniolepiota

are in an unre-

solved polytomy, sister to a well supported clade of

Barcheria

and a monophyletic

Agaricus

wariatodes

and

A. chartaceus

are sister taxa in a well

supported sect.

Minores

,and

A. eburneocanus

and

inapertus

are in a weakly supported sect.

Arvenses

but

with affinities to different species.

Agaricus inilleasper

in a weakly supported clade with

A. campestris

(sect.

Agaricus

)and

A. langei

and

A. silvaticus

(sect.

Sanguinolenti

Agaricus erythrosarx

is in a strongly

supported clade of taxa belonging to sect.

Sanguino-

lenti

.Finally

A. melanosporus

is sister to all remaining

species of

Agaricus

but without strong support or

affinities to any previously described section of

Agaricus

Macrolepiota/Chlorophyllum ITS dataset.

Species of

Macrolepiota

are grouped in a clade with a high PP (0.97).

Both

Micropsalliota

(PP 1.0) and

Leucoagaricus

(PP 1.0)

form a clade sister to a paraphyletic

Chlorophyllum

and a

strongly supported (PP 1.0) polytomy that groups

Barcheria

and

Agaricus

with

Heinemannomyces

Clar-

keinda

Coniolepiota

and

Eriocybe

(FIG.2).

Multiple accessions of

Chlorophyllum agaricoides

clustered together but some genetic heterogeneity is

apparent (F

IG. 2). Three subclades were recovered

within

Macrolepiota

matching currently recognized

sections

Volvatae

Macrosporae

and

Macrolepiota

(Vel-

linga 2003, Ge et al. 2010). Two of the new Australian

sequestrate species,

M. turbinata

and

M. vinaceofibril-

losa

, grouped within sect.

Macrolepiota

but do not form a

monophyletic group. The position of the third species,

M. gasteroidea

, is currently unresolved but morpholog-

ically appears to be related to this same group of taxa.

Calibrated phylogeny: the evolutionary origin of the

sequestrate sporocarp form.—

Based on a divergence

time between

Agaricus

and

Chlorophyllum

73 000 000 y (F

IG.3, SUPPLEMENTARY FIG.2),the

following divergence times are estimated (million

years ago 5 Ma):

Macrolepiota

Chlorophyllum

Agaricus

: 83 Ma [95% CI 72–98];

Barcheria

:65Ma

[95% CI 57–72];

Agaricus

: 34 Ma [95% CI 26–43].

Substitution rates were 0.0012–0.0101 substitutions

per site per million years. Character mapping based

on maximum likelihood (one rate) reconstructed

most nodes as the agaricoid form; the only exceptions

being the ancestors to clades of solely sequestrate

forms 2

Agaricus chartaceus

A. wariatodes

;

Barch-

eria

; and

Macrolepiota turbinata

(matrix in SUPPLE-

MENTARY TABLE

II). Width of branches are displayed

as thick or thin, according to the most likely recon-

structed state. A pattern seen in this character mapp-

ing is that sequestrate forms in these three lineages

tend to have evolved recently and since the mid-

Miocene (, 15–0 Ma). Stem lineages of

Agaricus

melanosporus

and

Barcheria

date back approximately

, 34 Ma and , 65 Ma respectively.

TAXONOMY

Barcheria willisiana

T. Lebel, in Lebel, Thompson &

Udovicic, Mycol. Res. 108(2):211 (2004)

IG.4A,B

Additional specimens examined

: WESTERN AUSTRALIA.

Kellerberrin, Site 17B, Higginson Road, 16 km along

Bencubbin-Kellerberrin Road, 29 Jun 1998,

I.C. Tommerup

H7518

[PERTH 07626886]; north of Kellerberrin, Site 17

(near Tree 22), Higginson Road, 16 km along Bencubbin-

Kellerberrin Road, 5 Sep 1995,

I.C. Tommerup H7306

[PERTH 07584407]; Corrigin Shire, Grylls Road, Valem

Farm, Paddock 12 (Todd’s Lounge), 13 Aug 1999,

Catchpole & S. Bolsen broek H7564

[PERTH].

498 MYCOLOGIA

Notes

Barcheria willisiana

is known only from a few

sites within drier woodlands of Western Australia,

associated with

Eucalyptus wandoo

E. salmonophloia

and

E. kondineninsis

. Additional specimens cited here

are of material collected more than 500 km west of the

type locality.

Barcheria

is supported as a sister taxon to

a monophyletic

Agaricus

in analyses of the ITS (FIG.1)

and in a strongly supported clade with species of

Agaricus

but with no resolution of generic boundaries

in analyses of LSU (S

UPPLEMENTARY FIG. 1).

Barcheria

retained here as a distinct genus, due to the distinctive

appearance of the sporocarp, characters of the spores

and strong support in the ITS phylogeny.

Agaricus chartaceus

T. Lebel sp. nov. FIGS. 4C, 5

Mycobank MB519924

Etymology: (L). papery 5 chartaceus, in reference to the

papery texture of dry sporocarps.

FIG. 1. Bayesian phylogram of

Agaricus

based on ITS data. Thick branches indicate support by posterior probabilities of

. 0.95. All sequestrate taxa are in boldface.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

499

Sporocarpum 12–38 mm altum, 12–27 mm diametro,

subglobosum vel elongatum apice convexo, albidum vel

cremeum, sericeum laevigatum, nitidum; contextus

non mutabilis. Hymenophorum pallido-flavidum dein

fumeum, ad initium minute labyrinthium-loculatum,

cellulis dein pulverescentibus. Stipes uterque basalis 8–

10 3 2–5 mm vel robustus percurrens 18–42 3 2–6 mm,

cremeus vel pallido-cinereus. Odor curcuma simulans.

Sporae 7.0–8.0 3 5.0–6.0(–7.0)

mm, subglobosae vel

obovatae, valde dextrinoideae.

Holotypus

: Western Australia. 86.2 km E of Broome

along the Great Northern Highway, 10 Feb 1993,

N.L.

Bougher H6271

[PERTH07582757].

Sporocarps hypogeous to emergent, gregarious to

solitary. Pileus 12–38 mm high 3 12–27 mm diam,

irregularly subglobose to elongate, apex convex,

white to pale cream, silky smooth, shiny, thin, papery

when dry, sometimes finely cracked; margin fused to

stipe by veil remnants, concolorous with pileus.

Context fleshy, thick, white to cream, not changing

color. Hymenophore pale yellowish becoming grayish

tan but not darkening further, initially minutely

labyrinthine-loculate; becoming powdery. Stipe either

an inserted basal pad, 8–10 3 2–5 mm, or robust,

percurrent, 18–42 3 2–6 mm diam, projecting well

beyond margin with slightly bulbous base, cream to

pale grayish cream, fibrous; context cream, no color

change. Odor of curry; flavor mild. Spores 7.0–8.0 3

5.0–6.0(–7.0)

mm, mean 7.2 3 5.9 mm, Q 5 1.0–1.3,

subglobose to obvate, mostly symmetric, pale yellow-

brown in KOH, dextrinoid. Basidia 7–16 3 8–12

mm,

clavate to broadly clavate; sterigmata two. Pileipellis a

cutis, 5–35

mm wide, of compact subparallel hyaline

hyphae 2–5

mm diam, overlying a broad context, 70–

250

mm wide, of slightly inflated hyphae 5–8 mm diam.

Clamps absent in all tissues.

Additional specimens examined

: WESTERN AUSTRALIA.

Gibson Desert, Patience Well, 6 Jun 2001,

C.P. Campbell

[PERTH05848261]; Coolgardie, Newman Rocks 50 km W of

Balladonia, 1 km N of Eyre Highway, 27 May 2003,

K. Symes

FIG. 2. Bayesian phylogram of

Chlorophyllum

and

Macrolepiota

based on ITS data. Thick branches indicate support by

posterior probabilities of . 0.95. All sequestrate taxa are in boldface.

500 MYCOLOGIA

1234/03

[MEL2231690]. SOUTH AUSTRALIA. Flinders

Ranges, Oraparinna Station, North of Wilpena Pound near

Blinman, 3 Sep 2000,

Catcheside 758

[AD]. NORTHERN

TERRITORY. 61 km N of Alice Springs, 28 Jun 1968,

H.A.

Morrison 6

[MEL202451]; 15 miles W of Ayers Rock, 30 Jun

1967,

A.C. Beauglehole 22929

[MEL260729]; George Gill

Range, west of Farrers Spring, 26 miles NW of Wallara

Ranch, 3 Jul 1967,

A.C. Beaglehole

[MEL260703].

Distribution and habitat

: Found in the drier regions

of central Western Australia, South Australia and

northern Australia. Associated with

Allocasuarina

and

Callitris

woodland.

Notes

Agaricus chartaceus

resembles

A. wariatodes

both macro- and microscopically, however the pileus

is papery not silky when dry, the fragrence is of curry

FIG. 3. Calibrated phylogeny of Agaricaceae species based on ITS data. The maximum clade credibility tree with posterior

probabilities is shown. Character trace of sequestrate form using a one-rate model under maximum likelihood. The best

supported state on each branch is represented by width; sequestrate lineages are shown by wide/bold branches and agaricoid

lineages by narrow branches.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

501

rather than marzipan, and the basidia are two- rather

than four-spored. Affinities lie with a strongly

supported sect.

Minores

(FIG. 1).

Agaricus eburneocanus

T. Lebel sp. nov. FIG. 4D, 6

MycoBank MB519925

Etymology

: (L) cream or ivory 5

eburneus

, gray 5

canus

(light) in reference to the sporocarp pellis and stipe.

Sporocarpum 15–20 mm altum, 25–28 mm diam,

late convexum apice plano, ad centrum eburneum vel

in margine late pallido-cinereum, margo tenuis

appendiculatus; contextus non mutabilis. Hymeno-

phorum brunneum trama argento-cinerea, ad initium

minute loculata vel labyrinthina, cellulis pulverescen-

tibus. Stipes 25–30 mm altus, 5–10 mm diam,

percurrens, tomentosus, cinereo-cremeus basi luteo;

contextus cremeus dein armeniacus. Sporae 8.0–

13.5(–16) 3 6.0–8.0

mm, variabiles amygdalinae vel

angulato-ovoideaea, plerumque elongatae ellipsoi-

deae vel ovoideae, infirme dextrinoideae.

Holotypus

: Western Australia. Site 17b, Higginson

Road (north), 16 km along Bencubbin-Kellerberin

Road, 29 Jun 1998,

I.C. Tommerup and S. Bolsenbroek

H7528

[PERTH07626789].

Sporocarp solitary to gregarious, hypogeous to

emergent. Pileus 15–20 mm high 3 25–28 mm diam,

broadly convex with convex to plane apex, white to

cream centrally to pale gray in broad marginal zone,

overlain by fine brown fibrils; margin initially

attached to stipe by cream partial veil, becoming free

in small sections as pileus expands. Context not

changing color. Hymenophore dark rich brown with

white to silvery gray trama, loculate to labyrinthiform

with minute chambers; becoming pulverant. Stipe-

columella 25–30 mm long 3 5–10 mm diam,

percurrent, cylindrical, appressed matted fibrillose,

white to cream with grayish tinge, with yellowish stains

basally; context white to cream becoming gradually

pale apricot. Odor not distinctive. Spores 8.0–13.5

(–16) 3 6.0–8.0

mm, mean 5 10.2 3 6.8 mmQ5 1.3–

1.7, variable shape, amygdaliform or angular-ovoid,

predominantly long ellipsoid or ovoid, brown in

KOH, weakly dextrinoid. Basidia 35.5–45.0 3 16.0–

20.5

mm, broadly clavate; sterigmata short, four rarely

FIG. 4. Sporocarps of taxa. A.

Barcheria willisiana

(MEL2151446; photo B. Archer). B.

Barcheria willisiana

(H7518; photo

N. Bougher). C.

Agaricus chartaceus

(holotype H6271; photo T. Lebel). D.

Agaricus eburneocanus

(photo N. Bougher; holotype

H7528). E.

Agaricus erythrosarx

(photo W. Dunstan MURU6080).

502 MYCOLOGIA

FIG.5.

Agaricus chartaceus

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and subhymenium, D. spores.

Bar 5 10

mm.

FIG.6.

Agaricus eburneocanus

. Illustrations of A. sporocarps, B. basidia and subhymenium, C. spores. Bar 5 10 mm.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

503

two. Pileipellis a cutis of hyaline interwoven hyphae 3–

mm diam, overlying a broad context 100–230 mm

wide, of hyaline regular to somewhat interwoven

hyphae 3–8

mm diam and abundant inflated elements

15–47 3 5–26

mm. Clamps absent from all tissues.

Additional specimens examined

: WESTERN AUSTRALIA.

Seven km NE of Dowerin, approximately 200 m SW of refuse

dump at AMERY, 11 Jul 2002,

W.A. Dunstan

[MURU5927].

Distribution and habitat

:Foundin

Eucalyptus

wandoo

and

E. salmonophloia

woodland, in leaf litter

at the base of trees.

Notes

Agaricus eburneocanus

is characterized by the

agaricoid sporocarps, dark brown hymenophore with

silvery gray trama, and grayish cap. Affinities lie within

a weakly supported sect

Arvenses

, in a subclade with

an undescribed Australian epigeous species,

Agaricus

sp. ‘‘B’’ (FIG. 1).

Agaricus erythrosarx

T.Lebel sp. nov. FIGS. 4E, 7

Mycobank MB519926

Etymology

: (Gk)

erythrosarx

5 red flesh, for the reddening

reaction of the flesh when cut or exposed.

Sporocarpum (20–)30–60 mm altum, (40–)65–

120 mm diam, convexum vel plano-convexum, apice

depresso initio ad maturationem ellipsoideum irre-

gulariter apice plano vel depresso, albidum vel

cremeum, nitidum sericeum radiale fibrillosum,

superato squamato vinaceo sparso, velum partiale,

albidum sericeum; contextus albidus mutabilis

erubescens. Hymenophorum sublamellatum, cellulis

grandis cassis labyrinthinis compostium, pallido-hin-

nuleum dein brunneum, in siccitate leviter pulver-

atum trama cremeum dein aurantiaco-ruber. Stipes

(24–)30–60(–80) mm altus, 15–25 mm diam, percur-

rens cremeus; contextus cremeus tarde rubescens.

Sporae 7.0–9.0 3 5.0–7.0

mm, subglobosae vel late

ellipsoideae, non dextrinoideae.

Holotypus

: Western Australia. Elyra Brook, Bolgart

(off Blood Road), 25 Jul 2004, W.A. Dunstan

[MURU6080].

Sporocarp solitary or gregarious, hypogeous to

emergent. Pileus (20–)30–60 mm high 3 (40–)65–

120 mm diam, convex to planoconvex, apically

depressed when young, usually irregularly ellipsoid

with flat apex or slightly depressed at center when

mature, white to cream, shiny silky radially fibrillose,

overlain with scattered vinaceous fibrils that may

aggregateintofinescales,moreabundantnear

center; margin remaining attached to stipe by veil of

dense white silky fibrils. Context fleshy, compact, 2–

6 mm thick, white, reddening when cut. Hymenophore

pale faun maturing to dark brown, trama cream but

slowly and irregularly reddening (to orange-red) when

exposed, sublamellate to labyrinthine, of large, disor-

ganized empty chambers; moist initially becoming

slightly pulverant. Stipe (25–)30–60(–80) mm long 3

15–25 mm diam, percurrent, cylindrical or tapering

slightly toward base, solid initially often becoming

FIG.7.

Agaricus erythrosarx

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and hymenophoral trama,

D. spores. Bar 5 10

mm.

504 MYCOLOGIA

slightly hollow at maturity, cream with reddish tints,

with scattered vinaceous fibrils in lower section;

context cream then slowly reddening, smooth, dry.

Odor and flavor mildly spicy. Spores 7.0–9.0 3 5.0–

7.0

mm, mean 8.1 3 6.3 mm, Q 5 1.1–1.4, subglobose to

broadly ellipsoid, slightly asymmetric, brown in KOH,

not dextrinoid. Basidia 20.0–28.5 3 9.5–14.0

mm,

clavate; four-spored. Pileipellis a cutis, 45–110

mm wide,

of hyaline, interwoven to periclinal hyphae, 5–10

diam. Clamps absent in all tissues.

Additional specimens examined

: WESTERN AUSTRALIA.

Paddock 53 (remnant opposite farmhouse), Grylls Road,

Valerma Farm, west of Corrigin, 19 Jul 2000,

N. Bougher

Tommerup

R. Campbell H7638

[PERTH].

Distribution and habitat

: Found at the base of trees

Eucalyptus eremophila

and

E. loxophleba

subsp

loxophleba

woodland.

Notes

Agaricus erythro sarx

resembles

A. melanos-

porus

both macro- and microscopically. Both species

have large pale sporocarps with scattered vinaceous

fibrils on the surface, a sublamellate hymenophore

that becomes pulverant, reddening pileal flesh and

small spores. Differences include the red-orange

staining reaction of the hymenophoral trama and

reddening of the stipe context in

A. erythrosarx

and

lack of color changes in

A. melanosporus

(some dingy

brown stains at base of stipe), and narrower basidia

(5–9

mm vs. 9.5–14 mm) and slightly larger spores in

A. melanosporus

Agaricus erythrosarx

has strong

affinities to section

Sanguinolenti

, including an

undescribed agaricoid Australian taxon,

Agaricus

sp. ‘‘C’’ (FIG.1).

Agaricus inilleasper

T. Lebel sp.nov. FIGS. 8, 10A

Mycobank MB519927

Etymology

: (L)

5 in,

ille

5 the (usually not used),

asper

5 rough, is in reference to the location of the type

collection on a golf course.

Sporocarpum 25 mm altum, 30 mm diam, con-

vexum, albidum vel pallido-cremeum maculatum

ferrugineum et lutescens, laevigatum vel areolatum;

contextus non mutabilis. Hymenophorum fumeum,

sublamellatum vel labyrintho-loculatum irregulariter,

pulverescens. Stipes 20 mm altus, 5 mm diam,

percurrens, albidus; contextus pallido-cremeus.

Sporae 6.0–8.0 3 5.0–6.0 (–7.0)

mm, ellipsoideae vel

subglobosae, infirme dextrinoideae.

Holotypus

: Queensland. Townsville, Willows golf

course, fairway 9th hole, 9 Apr 1989,

M. Castellano

H4452

(holotype PERTH; isotype DAR).

Sporocarp solitary, emergent. Pileus 25 mm high 3

30 mm diam, convex, white to pale cream with some

tan patches, staining yellowish where handled,

FIG.8.

Agaricus inilleasper

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and subhymenium, D. spores.

Bar 5 10

mm.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

505

smooth or obscurely wrinkled or areolate; margin

remaining fused to stipe. Context 1–3 mm thick, firm,

white in section with no color change. Hymenophore

brownish gray, sublamellate to irregularly labyrin-

thine-loculate; becoming powdery. Stipe 20 mm long

3 5 mm diam, percurrent, with some small chambers,

white; context pale cream, no staining reactions.

Odor and flavor not recorded. Spores 6.0–8.0 3 5.0–

6.0(–7.0)

mm, mean 7.3 3 5.6 mm, Q 5 1.2–1.4,

ellipsoid to subglobose, sometimes obvate, symmetri-

cal, hyaline initially then pale brown in KOH, weakly

dextrinoid. Basidia 22–29 3 6–8

mm, cylindrical to

narrowly clavate; sterigmata four, rarely two. Pileipel-

lis a narrow cutis of compact subparallel hyaline

hyphae 2–7

mm diam, overlying a context of hyaline

hyphae 3–8

mm diam and irregular inflated elements

3–42 3 7–28

mm. Clamps absent in all tissues.

Distribution and habitat

: Known only from the

holotype collection, from northern Queensland.

Found among grass.

Note

Agaricus inilleasper

may be distinguished from

other species by the pale sporocarp staining yellowish,

brownish gray sublamellate to irregularly labyrinthine

hymenophore, small spores. Although it is preferable

to not describe taxa from single collections (known

only from the type), this species is distinctive. At first

glance it resembles

Agaricus bisporus

in sect.

Duploan-

nulati

, however analysis of ITS places it in a weakly

supported clade with some species from sect.

Sanguinolenti

and sect.

Agaricus

FIG.9.

Agaricus melanosporus

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. spores. Bar 5 10 mm.

506 MYCOLOGIA

Agaricus melanosporus

(Berk.) T. Lebel comb. nov.

IGS. 9, 10B, C

Mycobank MB297191

T: Western Australia. Swan River,

Drummond 180

(K;

BPI704933)

;

Endoptychum melanosporum

(Berk.) Singer & A.H. Sm.,

Brittonia

10:220 (1958)

;

Secotium melanosporum

Berk.,

London J. Bot

. 4:62

(1845)

Endoptychum kolya

Grgur.,

Larger Fungi of South

Australia

437 (1997). T: South Australia: 7 Jul 1923,

Kinchina,

JBCleland

[AD9767]

Additional illustration s

: Griffiths (1985) A field guide to

the larger fungi of the Darling Scarp and south west of

Western Australia. p 59 (figures lower right).

Sporocarps solitary to gregarious, hypogeous to

emergent. Pileus (18–)25–45 mm high 3 (20–) 35–

72 mm diam, subglobose to turbinate, planoconvex

FIG. 10. Sporocarps of taxa. A.

Agaricus inilleasper

(Holotype H4452). B.

Agaricus melanosporus

(Type K,

Secotium

melanosporum

, Swan River 180). C.

Agaricus melanosporus

(Corda

Icones fungorum

, Pl VI fig. 19–24). D.

Agaricus wariatodes

(Type K,

Secotium coarctatum

, Swan River 181). E.

Agaricus wariatodes

(Corda

Icones fungorum

, Pl VI fig. 25–30). F.

Agaricus

wariatodes

(TWM 1589; photo TW May).

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

507

with somewhat squarish edges, sometimes with

depressed center, pale tan to cinnamon drab or

warm brown, smooth to very finely fibrillose with

slightly darker fibrils scattered over surface; margin

incurved to stipe with fibrillose veil remnants

appendiculate, pale concolorous with pileus, en-

closing hymenophore. Context fleshy, white turning

pale brownish with pale reddish tint on exposure.

Hymenophore initially pallid olive brown then dark

brown to purple brown, sublamellate to labyrin-

thine loculate; becoming slightly powdery. Stipe

(35–)50–120 mm long 3 6–16(–21) diam, percur-

rent, robust, equal, pale tan with dingy brown tints,

particularly toward base, silky. Odor and flavor not

recorded. Spores 8.0–10.0(–11) 3 6–7.5(–8)

mm,

mean 8.9 3 6.8

mm, Q 5 1.2–1.4, subglobose to

broadly ellipsoid or obovate, many asymmetric, pale

yellowish brown to brown in KOH, weakly dextri-

noid. Basidia 17–28 3 5–9

mm, cylindrical to clavate;

sterigmata four. Pileipellis a cutis of subparallel,

interwoven, hyaline hyphae 2–7

mmdiamand

scattered inflated elements 10–18 3 7–12

mm,

overlying a broad context, 170–350

mmwide,of

loosely interwoven hyaline hyphae, 3–6

mmdiam.

Clamps at base of basidia, but not observed in pileal

or hymenophoral trama.

Specimens examined

: SOUTH AUSTRALIA.

J.B. Cleland

[AD9763]; Monarto South, Sep 1920,

J.B. Cleland

[AD2971

5 AD48433; MELU6252F]; region 9, Murray, Halidon,

J.B.

Cleland

[AD22633];

J.B. Cleland

[AD9765]; Monarto South,

27 May 1921,

J.B. Cleland

[AD20223; PDD8396; looks like

photo in Cunningham]; Karoonda, 30 Jun 1946,

J.B.Cleland

[AD22628]; Kinchina, Adelaide, 8 Jun 1925,

J.B. Cleland

[AD22621; PDD8388];

J.B. Cleland

[BPI718210; CG. Lloyd

#734]. VICTORIA. Mallee Research Station, Walpeup,

Jun 1955,

E.M. Packe & A. Mann

[MELU6253F]; 4 km

from Inglewood, Green Hills, 25 May 1988,

F. Watts

[MEL2032783]. WESTERN AUSTRALIA. Toodyay, 20 Aug

1983,

K. Griffiths

[PERTH00962007].

Material held in US National Fungus Collections

(BPI): AUSTRALIA. BPI704932 (packet labeled ex-

Massee; fragments only), BPI704934 (packet labeled

ex-Paris; fragments only).

Distribution and habitat

: Found in open disturbed

ground and compacted rocky soil.

Note

Agaricus melanosporus

is characterized by the

large sporocarps with dark brown hymenophore at

maturity and broadly ellipsoid to obovate spores. The

only difference between earlier descriptions in

Berkeley (1845) and Cunningham (1935) was the

shape of the spores—obovate to elliptical in the

former rather than subglobose to ovate in the latter.

No observable morphological differences were found

on comparison of type material of

S. melanosporum

(Drummond 180) and

E. kolya

(AD9767) or other

more recent collections. Although no sequence was

obtained from type material of

S. melanosporum

(K),

the collections examined here fit nicely with Berke-

ley’s original concept and the illustration of type

material in Corda (1854) (F

IG. 10C).

Agaricus melanosporus

lacks a spore hilar cap and

has a cuticular pileipellis, characters of

Agaricus

rather than other lineages within the Agaricaceae.

Analysis of molecular data places this taxon at the

base of a strongly supported

Agaricus

, but relation-

ship to a particular section remains unresolved.

This species is a priority for recollection because the

most recent specimens are more than 25 y old (1983).

New material would be useful to add morphological

characters and also to obtain more sequence data.

Agaricus pachydermus

T. Lebel sp. nov. FIG.11

Mycobank MB519928

Etymology:

(L)

pachy

5 thick; -

derma

5 skin, in reference

to the thick pellis.

Sporocarpum 15–28 mm altum, 10–23(–28) mm

diam, subglosum apice planoconvexum, pallido-cre-

meum vel pallido-brunneum, laevigatum vel subtiliter

fibrillosum siccum; contextus non mutabilis. Hyme-

nophorum pallido-brunneum vel purpureo-brun-

neum, sublamellatum vel labyrinthinum, pulveres-

cens. Stipes 5–16 mm altus, 3–6 mm diam, percurrens

vel truncatus, cremeus; contextus cremeus non

mutabilis. Sporae (6–)7–8 3 (5–)6–7

mm, globosae

vel subglobosae, non dextrinoideae.

Holotypus

: South Australia. Flinders Ranges, within

Wilpena Pound (ca. 40 km NE of Hawker), 21 Sep

1971,

G.A. Crichton P11

[MEL2059965].

Sporocarps emergent, solitary to gregarious. Pileus

15–28 mm high 3 10–23(–28) mm diam, subglobose

to planoconvex, light cream to pale brown, smooth to

finely fibrillose, becoming brittle when dried. Context

fleshy when fresh, compact, white, no color change.

Hymenophore pale brown to dark purple-brown,

sublamellate to elongated labyrinthine; moist becom-

ing powdery. Stipe 5–16 mm long 3 3–6 mm diam,

percurrent or truncated, pale cream, fibrous, context

pale with no staining reactions, protruding shortly

below excavated hymenophore. Odor not recorded.

Spores (6–)7–8 3 (5–)6–7

mm, mean 7.4 3 6.7 mm,

Q 5 1.0–1.3, globose to subglobose, some slightly

asymmetric, pale yellowish brown to brown in KOH,

non-dextrinoid. Basidia 15–22 3 5–9

mm, elongate

clavate to cylindrical; sterigmata four. Pileipellis a

cutis, 24–42

mm wide, of subparallel, interwoven,

hyaline hyphae 2–5

mm diam and scattered inflated

elements 7–14 3 5–10

mm, overlying a context, 45–

mm wide, of loosely interwoven hyaline hyphae, 2–

mm diam. Clamps not observed.

Specimens examined

: NORTHERN TERRITORY. Raggatts

Creek, Glen Helen Station west of Mount Zeil, 22 Jul 1966,

508 MYCOLOGIA

J.H. Willis

[MEL269764]. NEW SOUTH WALES. Southern

Tablelands, paddock in Yass District, on the way to fossil

banks, 22 May 1968,

G.A. Crichton L39

[MEL2058665; K;

DAR 26446]; Haddens Corner, Yass, 20 May 1968,

G.A.

Crichton L65

[MEL 2058669]. WESTERN AUSTRALIA.

Carnarvon, 18 May 1971,

A.M. Young 134

[BRIP24660].

Distribution and habitat

: Found in arid regions, in

open grassy areas or claypans in flat mulga country in

NT, NSW, SA and WA.

Notes

Agaricus pachydermus

is similar to

A. war-

iatodes

and

A. chartaceus

, however the overall sporo-

carp shape differs, being more agaricoid. The pileus is

brittle and fragile when dried, often cracking to expose

the powdery spore mass, the hymenophore is dark

brown to purplish brown fresh versus pale tan to ocher,

lacks strong odors, and the spores are non-dextrinoid.

Unfortunately it proved difficult to obtain sequences

from any of the collections examined. Affinities of this

taxon to sections within

Agaricus

remain unresolved.

Agaricus wariatodes

(Grgur.) T. Lebel comb. nov.

IGS. 10D–F, 12

Mycobank MB443370

T: Western Australia. Swan River, Drummond 181 K;

BPI704922.

;

Endoptychum wariatodes

Grgur., Larger Fungi of South

Australia 436 (1997). T: South Australia. Adelaide,

Eagle on Hill, 26 Apr 1943,

J.B. Cleland

[AD9773].

Secotium coarctatum

Berk.,

London J. Bot

. 4:63 (1845).

non

Agaricus coarctatus

Cooke & Massee in Cooke

Grevillea

18:2 (1889).

Endoptychum moongum

Grgur.,

The larger fungi of South

Australia

435 (1997) T: South Australia. Greenhill

Road, 9 Apr 1932,

J.B.Cleland

[AD9759].

Nom

dubium Gautieria tasmanica

Rodway

Pap. & Proc.

Roy. Soc. Tasmania

1928:72 (1929). T: Tas. Rodway. non

G.Cunn. (1938); non

Agaricus tasmanicus

Berk. In Hook. F.

Fl. Tasman

. 2:245 (1859).

Additional illustrations

: McCann (2003 as gastroid

Agar-

icus

)

Australian fungi illustrated

p 69 (top left).

Sporocarps hypogeous to emergent, gregarious to

solitary. Pileus 8–18(–35) mm high 3 8–23(–34) mm

diam, irregularly subglobose to elongate or turbinate,

apex convex to planoconvex, cream to pinkish buff,

especially in creases, rarely pale dingy brown, silky

smooth to finely fibrillose, rarely drying faintly

tessellated; margin generally fused to stipe, veil

remnants appendiculate on some sporocarps, cream,

persistent. Context fleshy, firm texture, white to

cream, not changing color. Hymenophore initially

pinkish buff to yellowish buff, maturing to grayish

brown or rich olive-brown, minutely labyrinthine-

loculate; moist initially, becoming powdery. Stipe

(8–)17–40 mm long 3 2–6 mm diam, percurrent,

either barely projecting beyond margin with slightly

bulbous base or long, well developed, equal, cream to

pale grayish cream, fibrillose; context cream, no

bruising reactions. Odor weak to strong marzipan,

less fragrant in younger specimens, apparent even

after drying; flavor mild. Spores (5.5)7.0–8.0(9.0) 3

(5)6.0–7.5(8.5)

mm, mean 7.2 3 6.4 mm, Q 5 1.0–1.3,

globose to subglobose or sometimes obvate, mostly

FIG. 11.

Agaricus pachydermus

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and subhymenium, D. spores.

Bar 5 10

mm.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

509

symmetric, hyaline initially then pale yellow-brown to

pale brown in KOH, dextrinoid. Basidia (8)11–18 3

7–15

mm, clavate to broadly clavate, sterigmata two or

four. Pileipellis a cutis, 5–25

mm wide, of compact

subparallel hyphae 2–5

mm diam, hyaline to pale

yellowish in KOH, overlying a broad context of

slightly inflated hyphae 5–12

mm diam. Clamps absent

in all tissues.

Specimens examined

: VICTORIA. Wimmera, Vectis, Quan-

tong Cemetary Road, B. Maraoske property, 10 Jun 2003,

TWM 1589 [MEL2326250]; Grampians, Mount Arapiles,

road to summit, 1.6 km from Wimmera Highway, 8 Jun

2003, TWM 1584 [MEL2326251]; Arnold West, 1 Jun 1938,

J.H.Willis [MEL1052387]; near Tarnagulla, Arnold West,

Apr 1938, C.L. Barrett [MEL1052366]; 12 km SE of Stawell,

6 Jun 1987, I.R. McCann GAC B109 [MEL2058458,

MEL2292966]; about 12 km SE of Stawell, 10 Jun 1987,

I.R. McCann [MEL2292963; K]; Stawell, in Three Jacks

Wildflower Reserve, 15 Apr 2001, I.R. McCann & T.H. Argyll

[MEL2292032]; near junction of Germanio and Mosquito

Creek roads, N of Stawell, 31 Mar 2001, I.R. McCann & T.H.

Argyll [MEL2292031, MEL2292033]; Arnold West near

Rheola, May 1939, H. Watts [MELU6249F]; Near Ingle-

wood, Green Hills, 13 Apr 1983, F. Watts [MEL2032782];

Riverina, Mitiamo Cemetary, Terrick Terrick State Park, 25

Apr 1996, J. Eichler 16 [MEL2096554]; Riverina, Boort area,

P. Weavers Road approx. 4.5 km SSW of Boort, 5 Apr 1999,

J. Eichler 109 [MEL2096555]; far NW District, Meridian

Road ca. 20 miles SW of Red Cliffs, 1 Aug 1968, J.H. Willis

[MEL695342]; 3.2 km north of Deep Lead, near Ararat, 5,

16 May 1968, I.R. McCann 16 [MELU E0617F (L38), DAR

26447, MEL2058664]. SOUTH AUSTRALIA. Flinders Rang-

es, Wilpena Pound, 19 Jun 2000, Catcheside 540 [AD];

Flinders Ranges, Wilpena Pound near Quorn, 11 Aug 2000,

Catcheside 681 [AD]; Flinders Ranges, Wilpena pound, 19

Jun 2000, Catcheside 528 [AD]; Beaumont, near Adelaide,

17, 24 May 1925, J.B. Cleland [AD22620]; Beaumont,

Adelaide, 20 May 1923, J.B. Cleland [AD20224]; Kinchina,

8 Jun 1925, J.B. Cleland [AD20215]; Beaumont, Adelaide,

Apr 1935, J.B. Cleland [AD22622]; 13 km south of Mount

Davies, Aug 1962, D.E. Symon [AD15829]; WARI, May 1960,

J.H. Warcup [AD10572]; Waite Arboretum, 13 May 1953,

J.H. Warcup [WAITE 3353]; 1935, J.B. Cleland [AD9769];

Highfield, Adelaide, 10 Jan. 1934, J.B. Cleland [AD9771];

Adelaide, J.B. Cleland [MELU6250F]; Dry Creek, 25 Jul

1934, Prof Johnston [AD20208]. NEW SOUTH WALES.

Narrabri, 2 Jun 1919, J.B. Cleland [AD20217]; Sydney, J.B.

Cleland [BPI718213; CG. Lloyd #528]; Riverina, Savernake

Station, Lifeboat, 20.5 km E of Berrigan. SANDS site 6, 4

Jun 2000, J. Trappe 25186; Abbotsford Bridge over Murray

River, near Wentworth, 26 Aug 1955, J.H. Willis 157

[MEL1053064]; Riverina, Kilpa farm, 17.5 km SE of

Berrigan. SANDS site 5, 6 Nov 2000, J.M. Trappe 26601

[MEL2276058]; Trentham Station North of Mildura, 11

Aug 1986, I. McCann B108 & T.H. Argyll [MEL2292967];

Riverina Highway, W of SANDS Plot 23, 23 Jun 2003, J.M.

Trappe 28630. WESTERN AUSTRALIA. Dryandra State

Forest, Tomingley Road, 15 Jun 1999, M. Brundrett & T.

Lebel 43 [MEL2063323]; Dryandra State Forest, Tomingley

Road, 13 Aug 2000, K. Syme & M. Hart [MEL2219013];

Toodyay, 20 Aug 1983, K. Griffiths [PERTH963062];

FIG. 12.

Agaricus wariatodes

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and subhymenium, D. spores.

Bar 5 10

mm.

510 MYCOLOGIA

[BPI704923; packet labeled ex-Paris, small fragments only].

AUSTRALIA. Herb Hookeriana 1867 (K).

Distribution and habitat

: Found in arid areas among

moss, lichens, small herbaceous plants and scattered

litter, or in bare ground or fine gravel, in woodlands

Allocasuarina

sp.,

Callitris

sp.,

Eucalyptus leucox-

ylon

E. microcarpa

E. drummondii

E. tectifica

Acacia armata

Notes

Agaricus wariatodes

is characterized by the

subglobose spores, pallid color and minute cells of

the hymenophore, mostly four-spored basidia, and in

most collections, a pleasant, strong fragrence of

marzipan in fresh specimens (also noticeable in many

dried collections).

Agaricus wariatodes

is a widespread

species, found in dry woodlands in all states except

Queensland and Tasmania. Affinities lie within a

strongly supported sect.

Minores

with sister taxon

chartaceus

and two undescribed Australian taxa (

aff.

semotus

‘‘spH’’; and

Agaricus

sp. ‘‘D’’) (FIG. 1). It

is possible there are a couple of cryptic taxa because

of some variation in the ITS. However no morpho-

logical characters have been found to support

distinguishing varieties at this stage.

On examination of type material of

S. coarctatum

(Drummond 181, K & BPI) no observable difference

was found in spore shape to the type of

E. wariatodes

(AD9773) or more recent collections. The holotype

packet of

E. moongum

(AD9759) contains only pulver-

ized fragments, but the spore size range and shape and

note of strong odor are typical of

Agaricus wariatodes

paratype of

E. moongum

(AD22638) has an organized,

lamellate hymenophore, a membranous annulus on the

saccate stipe, and dark brown, asymmetric spores. All of

these characters point to an agaricoid rather than a

sequestrate form of

Agaricus

, and this collection

therefore is excluded from this taxon.

Although no DNA sequence was obtained from

type material of

S. coarctatum

(FIG. 10D), the collec-

tions examined here fit Berkeley’s original concept

and the illustration of type material in Corda (1854)

IG. 10E). Two collections of

Gautieria tasmanica

are

listed in HO; however one is a species of

Austrogau-

tieria

and the second collection could not be found.

Notes suggest both of these collections were made

after the type collection date (Chang and Kantvilas

1993). Therefore we are calling

G. tasmanica

nom.

dubium and not proposing formal synonymy with

wariatodes

. The specific epithets ‘‘

coarctatus

’’ and

‘‘

tasmanicus

’’ are preoccupied in the genus

Agaricus

and as such are not available for this species (Berkeley

1859, Cooke 1889). The next available names are

‘‘

wariatodes

’’ and ‘‘

moongum

’’, which were published

at the same time (Grgurinovic 1997). Accordingly we

have chosen

A. wariatodes

as the new combination for

this taxon (Art.11.5).

Macrolepiota gasteroidea

T. Lebel sp. nov.

IGS. 13, 14A

Mycobank MB519929

Etymology

: (L)

gasteroidea

5 sac-like, in reference to the

gasteroid nature of the sporocarp.

Sporocarpum (30) 35–45 mm altum, (25) 50–65 mm

diam, subglosum vel elongatum ad basim decrescens

apice planum vel convexum, cremeum vel pallido-

brunneum squamatae latae vel angustaeleviter fuscatae

in pagina dispersae; contextus non mutabilis. Hymeno-

phorum initio pallido-brunneum maturita atroluteo-

brunneum, sublamellatum, cellulis grandis elongatis

labyrinthinis, pulverescentibus compositis. Stipes 25–

37 mm altus, (9)16–18 mm diam, percurrens robustis

ad basim decrescens, cremeus; contextus cremeus non

mutabilis. Sporae 14.0–17.5(–19) 3 10.0–12.0(–13.0)

mm, ellipsoideae, dextrinoideae; poris germinationum

praesentis, operculis hili praesentis.

Holotypus

: Western Australia. Ludlow State Forest,

10 Jun 1980,

N. Malajczuk H0052

[PERTH].

Sporocarps solitary or gregarious, hypogeous to

emergent. Pileus (30)35–45 mm high 3 (25)50–

65 mm diam, subglobose to elongate, tapering slightly

toward base, convex or plane at apex, cream to pale

tan, with slightly darker, broad to narrow scales

scattered over surface, giving areolate appearance in

dried specimens; margin incurved to stipe abruptly or

gradually, remaining fused to stipe. Context fleshy,

compact, 2–5 mm wide, white with no change in

color. Hymenophore pale tan initially becoming dark

yellowish brown, sublamellateto labyrinthine, cham-

bers large, empty; becoming powdery. Stipe 25–37 mm

long 3 (9)16–18 mm diam (broken off in larger

speciman), percurrent, robust, tapering slightly to-

ward base, not bulbous, pale tan, smooth to finely

fibrillose, context white to pale tan. Odor and flavor

not recorded. Spores 14.0–17.5(–19) 3 10.0–12.0

(–13)

mm, mean 15.9 3 10.9, Q 5 1.3–1.6, ellipsoid,

asymmetric, yellowish brown in KOH, dextrinoid;

germ pore present, small hilar cap present. Basidia

12.0–22.5 3 7.5–13.0

mm, clavate to broadly clavate;

sterigmata two. Pileipellis a weak trichoderm of

cylindrical interwoven pale brown, thick-walled hy-

phae 35–155

mm long 3 4–10 mm diam with terminal

cells slightly inflated 14–37 3 6–10

mm, obtuse or

occasionally tapered, overlying a broad context of

hyaline, thin-walled hyphae, mostly 2–3.5

mm diam

with scattered inflated hyphae 5–11

mm diam. Clamps

present at base of basidia, not common.

Additional specimens examined

: WESTERN AUSTRALIA.

Coastal Plain, 14 Apr 1983,

N. Malajczuk H0326

[PERTH];

Leederville,

Stoward

[BPI704142; cited by C.G. Lloyd (1918)

and Cunningham (1979) as

S. acuminatum

Distribution and habitat

:Foundin

Eucalyptus

gomphocephala

woodland.

EBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

511

Note

Macrolepiota gasteroidea

resembles

M. vina-

ceofibrillosa

; the main macroscopic difference is the

lack of color change of the flesh and overlying

vinaceous fibrillose squamules on the pileus of

gastero idea

. Both are rare taxa associated with a

restricted species of eucalyptus,

E. gomphocephala

native to a small region of southern Western

Australia. Analysis of LSU sequences place this taxon

within an unsupported

Macrolepiota

(SUPPLEMENTARY

FIG

. 1).

Macrolepiota turbinata

T. Lebel sp. nov.

IGS. 14B, 15

Mycobank MB519930

Etymology

: (L)

turbinatus

5 turbinate; in reference to the

sporocarp shape.

Sporocarpum (30–)50–110 mm altum, (30–)50–

90 mm diam, elongatum conicum vel subglobulosum

apice convexo vel planoconvexo, basim decrescentem

abrupte vel gradatim, pallido-cremeobrunneum sicca-

tate argenteum interdum vinaceum, laevigatum vel

squamatum concolorum; contextus non mutabilis.

Hymenophorum initio albidum maturita brunneum,

sublamellatum, cellulis grandis elongatis labyrinthi-

nis, pulverescentibus compositum. Stipes (37–)48–

90 mm altus, (8–)20–35 mm diam, percurrens

robustis ad basim bulbosus vel turbinatus, cremeus;

contextus cremeus. Sporae (12–)14(–17) 3 (8–)10

(–11)

mm, late ellipsoideae vel obovatae, dextrinoi-

deae; poris germinationum praesentis, operculis hili

praesentis.

Holotypus

: Western Australia. Agroforestry site. 30

Sep 1981,

N. Malajczuk

H0219 [PERTH].

Sporocarps partly or almost wholly epigean, sessile,

with few basal rhizomorphs. Pileus (30–)50–110 mm

high 3 (30–)50–90 mm diam, elongate conical or

squat and spherical, sometimes turbinate when

young, apex convex or planoconvex, tapering abrupt-

ly or gradually to short stipe, pale creamy tan, drying

silvery white, with a purplish tint on some sporocarps,

smooth overall or with scattered to dense, fine

fibrillose scales, concolorous or slightly darker than

FIG. 13.

Macrolepiota gasteroidea

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and subhymenium,

D. spores. Bar 5 10

mm.

512 MYCOLOGIA

pellis; margin remaining attached to robust stipe,

enclosing hymenophore. Hymenophore off-white to

tan initially, becoming dingy grayish brown to dark

brown at maturity, sublamellate to labyrinthine;

becoming powdery. Stipe-columella (37–)48–90 3

(8–)20–35 mm diam, percurrent, bulbous at base,

tapering slightly or abruptly, pale cream, minute

fibrillose scales scattered over surface, context cream;

persistent fibrillose veil remnants, concolorous with

pellis or slightly paler. Odor and flavor sweet,

pleasant. Spores (12–)14(–17) 3 (8–)10(–11)

mm,

mean 14.3 3 9.9

mm, Q 5 1.2–1.6, broadly ellipsoid to

slightly obvate, some pedicillate, pale yellowish brown

to brown in KOH, dextrinoid; germ pore present,

distinct, hyaline cap present. Basidia 9.0–14.5 3 16.0–

24.5

mm diam, spherical or broadly clavate; sterigmata

two or four. Pileipellis a patchy trichoderm of

cylindrical interwoven, thick-walled hyphae 70–

200

mm long 3 3–7 mm diam with terminal cells

slightly inflated 14–30 3 5–8

mm, obtuse or occasion-

ally tapered, with scattered brown pigmentation,

overlying a broad context of hyaline, thin-walled

hyphae, mostly 2–3.5

mm diam with scattered inflated

hyphae 5–9

mm diam. Clamps present at base of

basidia, not common.

Additional spec imens examined

: WESTERN AUSTRALIA.

Ludlow State Forest near Busselton, 17 Jun 2002,

Bradshaw

[MURUL055]; Murdoch University Campus, 22

Apr 1982,

R. Watling 14576

; Margaret River, 15 Apr 1963,

L.E. Souef 27

[PERTH00952516]; Yanchep, Aug 1963,

Newhouse 3

[PERTH00953571]; Margaret River, 24 Jun

1962,

L.E. Souef 18

, [PERTH00952036]; Murdoch University

carpark, 1975,

R.N. Hilton

[PERTH00959359]; Margaret

River,414SussexStreet,2May1964,

L.E. Souef

[PERTH00954624]; Jurien Bay, Mountt Benia, 18 May

1971,

R.N. Hilton

[PERTH00956295]; Yanchep, Aug 1963,

O.K. Miller

[PERTH00952591]; Margaret River, 12 May 1963,

L.E. Souef 28

[PERTH00951951]; Margaret River, 15 Apr

1963,

L.E. Souef 27

[PERTH00952060]; Ashdale, Coomal-

bidgup, near Esperance, 7 Apr 1974,

G. Bowen

[PERTH00958832]; Guildford, May 1967,

T.C..Daniell 29

[PERTH00955159]; Edgewater, 27 Apr 1982,

M. Daams

[PERTH00960934]; Waneroo, Apr 1982,

J. Daams Y624

[BRIP25269]; Margaret River, 414 Sussex Street, 30 May

1964,

L.E. Souef

[PERTH00954136]; Margaret River, 17 Jun

1962,

L.E. Soeuf 31

[PERTH00951501]; Esperance, 7 May

1971,

M.S. Daniell 143

[PERTH00958387]; Esperance, 20

May 1972,

T.C. Daniell

[PERTH00957771];

W. Stewart

[BPI704148 under the name

Secotium acuminatum

Distribution and habitat

: Emergent in sandy soils

under leaf litter, in mixed open, mostly coastal,

Eucalypt forest, including

Eucalyptus gomphocephala

E. marginata

E. calophylla

and

Agonis flexuosa

Note

Macrolepiota turbinata

is distinguished by the

overall shape of the sporocarp, with its tapering base

FIG. 14. Sporocarps. A.

Macrolepiota gasteroidea

(photo N. Malajczuk; holotype H0052). B.

Macrolepiota turbinata

(photo

W. Dunstan MURUL055). C.

Macrolepiota vinaceofibrillosa

(photo N. Bougher 0546).

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

513

to the stipe, smooth pale pileus and dark brown

hymenophore. The species appears to be restricted to

Western Australia but is fairly widespread in the

southern coastal region. Affinities to sect.

Macro-

lepiota

are supported by ITS analyses, with the closest

taxa

M. dolichaula

and

M. detersa

(FIG. 2).

Macrolepiota vinaceofibrillosa

T. Lebel sp. nov.

IGS. 14C, 16

Mycobank MB519931

Etymology

: (L)

vinaceo

5 vinaceous; -

fibrillosa

5 fibrillose,

in reference to the vinaceous fibrils on the pileus.

Sporocarpum 65 mm altum, 45 mm diam, tur-

binatum vel elongatum apice convexum, pallido-

brunneum squamatae latae concolores in pagina

despersae; contextus albidus roseolescens. Hymeno-

phorum initio pallido-brunneum maturita brunneum,

sublamellatum, cellulis grandis elongatis labyrinthinis,

pulverescentibus compositum. Stipes 90 mm altus, 18–

26 mm diam, percurrens robustis ad basim bulbosus,

albidus vel pallido-brunneus; contextus albidus roseo-

lescens. Sporae 12.5–15.0 3 10.0–12.0

mm, ellipsoideae

vel late ellipsoideae, dextrinoideae; poris germinatio-

num praesentis, operculis hili praesentis.

Holotypus

: Western Australia. Bold Park, Floreat

Park, 2 Jun 1980,

N. Malajczuk H0047

[PERTH].

Sporocarp solitary, hypogeous to emergent.

Pileus 50–65(–95) mm high 3 45–60(–70) mm

diam, turbinate-elongate with convex apex and

tapering slightly toward base, pale tan with scat-

tered to abundant vinaceous, mainly appressed

fibrillose-agglutinated scales; margin remaining

fused to stipe. Context fleshy, 2–5 mm thick, white

turning pale pinkish brown. Hymenophore pale

greenishbrownbecomingdulltodarkbrown,

sublamellate to labyrinthine, chambers large; be-

coming powdery. Stipe 75–90 mm long 3 18–26

FIG. 15.

Macrolepiota turbinata

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and hymenophoral trama,

D. spores. Bar 5 10

mm.

514 MYCOLOGIA

(–32) mm diam, percurrent, slightly bulbous then

tapering slightly toward base, fleshy when fresh,

white to pale tan, protruding below pileus 20–

30 mm; context white turning pale pinkish brown.

Odor and flavor not recorded. Spores 12.5–15.0 3

10.0–12.0

mm, mean 14.0 3 10.6, Q 5 1.2–1.4,

ellipsoid to broadly ellipsoid-ovoid, variable shape

and size, asymmetrical, yellowish brown in KOH,

dextrinoid; small germ pore present, hilar cap

present. Basidia 12.0–14.5 3 20.5–28.0

mm, clavate

to cylindrical; sterigmata four. Pileipellis a patchy

trichoderm of cylindrical interwoven brown pig-

mented, thick-walled hyphae 32–190

mmlong3 3–

mm diam with terminal cells slightly inflated 14–37

3 6–10

mm, obtuse or occasionally tapered, overly-

ing a broad context of hyaline, thin-walled hyphae,

septate, mostly 2–3.5

mmdiamwithscattered

inflated hyphae 5–11

mm diam. Clamps present at

base of basidia, not common.

Additional specimen examined

: WESTERN AUSTRALIA.

City Beach, Bold Park, in front of Ecology Centre, 9 Jul

2009,

N. Bougher 00546

[PERTH].

Distribution and habitat

:Foundin

Eucalyptus

gomphocephala

woodland.

Note

Macrolepiota vinaceofibrillosa

resembles

gasteroidea

. Microscopically

M. vinaceofibrillosa

spores

are slightly shorter and more broadly ellipsoid than

those of

M. gasteroidea

, the pellis hyphae are slightly

narrower with longer terminal cells, and the basidia

have four rather than two sterigmata. Affinities with

sect.

Macrolepiota

are supported by both morphology

and DNA (F

IG. 2). Unlike

Macrolepiota procera

and

allies,

Macrolepiota vinaceofibrillosa

and sister taxa

dolichaula

M. turbinata

M. gastroidea

and

M. detersa

generally have a pileus with slightly inflated terminal

elements that are not thick-walled, clamp connections

at the base of basidia, and lack big patches of velar

material on the pileus.

EY TO AUSTRALIAN SEQUESTRATE

GARICUS

ARCHERIA

AND

ACROLEPIOTA

SPECIES

1a. Sporocarps astipitate, soft, globose; flesh staining

orange reddish ............

Barcheria willisiana

1b. Sporocarps stipitate, firm, subglobose, or turbi-

nate; flesh may or may not change color ....... 2

2a. Spores with a germ pore and hilar cap present;

pellis a trichodermium; clamp connections

present at the base of basidia . .......... 3.

FIG. 16.

Macrolepiota vinaceofibrillosa

. Illustrations of A. sporocarps, B. pileipellis hyphae, C. basidia and hymenophoral

trama, D. spores. Bar 5 10

mm.

LEBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

515

2b. Spores with or without a germ pore, but hilar

cap lacking; pellis a cutis; clamp connections

almost always absent at the base of basidia . . . 5

3a. Flesh staining pinkish; pileus covered in fine

vinaceous squamules . . .

Macrolepiota vinaceofibrillosa

3b. Flesh not changing color; pileus smooth or with

broad, flat concolorous squamules . . .......... 4

4a. Pileus scaly, areolate in dried specimens; basidia

two-spored; spores 14.0–17.5(–19) 3 10.0–12.0(–13)

mm, ellipsoid .............

Macrolepiota gasteroidea

4b. Pileus smooth; basidia four-spored; spores (12–)

14.0(–17) 3 (8–)10.0(–11)

mm, broadly ellipsoid

toslightlyobvate ........

Macrolepiota turbinata

5a. Sporocarps with a strong fragence of almonds,

marzipan or anise ........................ 6

5b. Sporocarps without odor or with a different odor . . . 7

6a. Pileus thin, remaining fleshy; basidia four-

spored ...............

Agaricus wariatodes

6b. Pileus thin, becoming papery; basidia two-

spored ...............

Agaricus chartaceus

7a. Sporocarps large, generally . 40 mm diam (up to

120mm) .............................. 8

7b. Sporocarps smaller, generally , 25 mm diam (up

to30mm) ............................. 9

8a. Hymenophoral trama staining reddish orange

and stipe-context reddening

Agaricus erythrosarx

8b. Hymenophoral trama and stipe-context not

changing color .......

Agaricus melanosporus

9a. Pileus or hymenophore staining yellow when

bruised ..................

Agaricus inilleasper

9b. Pileus or hymenophore not staining or becoming

reddish ............................... 10

10a. Spores globose to subglobose 6–8 3 6–7.5

mm;

stipe flesh not changing color

..........

...................

Agaricus pachydermus

10b. Spores predominantly ellipsoid or ovoid

8–13.5 (–16) 3 6–8

mm; stipe flesh yellowish

then apricot at base . . .

Agaricus eburneocanus

DISCUSSION

Several studies have shown that the sequestrate

sporocarp form has arisen multiple times within the

Agaricaceae, with much of the diversification occur-

ring in

Agaricus

(Hopple and Vilgalys 1999, Vellinga

et al. 2003, Vellinga 2004, Gube 2009). The species of

Macrolepiota

reported here are an indication that the

sequestrate sporocarp form is more widely distributed

than initially thought. The three novel taxa are

derived from within sect.

Macrolepiota

and all are

currently known only from southern Western Austra-

lia. Given the similarities in appearance both macro-

and microscopically, overlap in distributions, it is

possible that

Macrolepiota gastroidea

is a two-spored

variant of

M. vinaceofibrillosa

, although analysis of

DNA data is inconclusive (S

UPPLEMENTARY FIG.1,

IG. 2). The sister taxa to

M. turbinata

and

vinaceofibrillosa

appear to be

M. dolichaula

(Austra-

lian/Asian) and

M. deters

, a recently described species

from Japan and China (Ge et al. 2010), rather than

either of the other Australian species,

M. clelandii

M. eucharis

(FIG. 2). Both

M. dolichaula

and

detersa

have medium to large, whitish sporocarps with

a long stipe, however

M. dolichaula

has small uplifted

pale squamules while

M. detersa

has large patch-like

brownish squamules (Ge et al. 2010). Intriguingly

mapping the distributions of Australian

Macrolepiota

species shows no overlap (Western Australia) in

sequestrate and sister agaricoid species (northeastern

Australia), the most likely explanation for current

distributions and relationships are that related taxa

are now extinct and/or documentation of taxa is

incomplete.

As with the epigeous taxa, spores of sequestrate

Macrolepiota

tend to be on the larger side (8–20 mm

long) and taxa also tend to have fairly large and fleshy

sporocarps; although in the case of the sequestrate

taxa the hymenophore becomes pulverant at maturi-

ty. Clamps are present at the base of basidia, but not

common, in the sequestrate forms, but this is variable

in both

Macrolepiota

and

Chlorophyllum

. Instead the

presence of a hilar cap covering the germ pore and

trichoderm versus a hymeniderm pellis are consid-

ered good characters differentiating

Macrolepiota

from

Chlorophyllum

. Without these characters it

would be difficult to differentiate sequestrate

Macro-

lepiota

species from

Chlorophyllum agaricoides

solely

on macromorphology.

No Australian sequestrate species of

Chlorophyllum

were discovered in this study, and Australian speci-

mens of

C. agaricoides

cited in Cunningham (1979)

are of

Agaricus wariatodes

‘‘Green Hill Rd, SA 9 April

1932’’, and

Macrolepiota gasteroidea

‘‘Leederville, WA,

F.W. Stoward’’.

Chlorophyllum

is paraphyletic in

analyses of both LSU and ITS, however the ‘‘

molybdites

’’group (

C. globosum

C. neomastoideum

agaricoides

) is well supported (SUPPLEMENTARY FIG.1,

IG. 2). The strong molecular support for

C. agar-

icoides

as a distinct taxon and lack of morphological

differences between specimens from a broad geo-

graphic area including much of Europe and North

America suggests that this is a stable, widespread

species undergoing little selection pressure for

speciation or that there has been a fairly recent

dispersal of the species across continents.

Our results and others show that the majority of

taxa once placed in the genus

Endoptychum

belong in

other genera. The only species remaining,

arizonicum

, cannot be placed with any certainty.

Endoptychum arizonicum

has extremely thick-walled,

white to pale yellow spores that lack a germ pore, a

non-percurrent stipe, no color changes on exposure

to air and a pileus that is a cutis rather than a

516 M

YCOLOGIA

hymeniderm (Shear 1902, Singer and Smith 1958,

Moreno et al. 2007). These characters would tend to

point to a closer affinity to

Agaricus

and

Barcheria

rather than

Chlorophyllum

. Gube’s (2009) combined

phylogenetic analyses of ITS and LSU were inconclu-

sive, showing

E. arizonicum

to be sister to

Agaricus

and

Chlorophyllum

. Of the other taxa that are

apparently sister to

Chlorophyllum

Barcheria

and

Agaricus

, both

Coniolepiota

and

Eriocybe

also have

pale or white spores but

Eriocybe

spores display an

immediate and strong dextrinoid reaction that is

lacking in

E. arizonicum

Endoptychum arizonicum

sporocarps lack the dense velar covering of pileus and

stipe and strong odors but do have scattered clamp

connections in the hymenophoral trama and at the

base of basidia such as

Coniolepiota

and

Eriocybe

(Vellinga et al. 2011).

Endoptychum arizonicum

remains somewhat in limbo, being retained as a

species of

Endoptychum

until further data is available.

In our analyses

Barcheria willisiana

also appears as

sister to

Agaricus

rather than within as suggested by

Gube (2009). Morphologically there is little resem-

blance to

Endoptychum arizonicum

, although both

have gastroid sporocarps, with a thin, fragile pileus, a

cuticular pellis and spores lacking a germ pore.

The diversity of native

Agaricus

species in Australia

is poorly known (May et al. 2006); thus putative sister

taxa to sequestrate species may be missing from

analyses. Our analyses indicated that the nine

sequestrate species in

Agaricus

are a result of at least

six transitions from agaricoid to sequestrate form,

with the highest diversity apparent in sections

Arvenses

and

Minores

. In various analyses by other

authors

Agaricus deserticola

Longula texensis

]isin

sect.

Arvenses

, which includes

A. eburneocanus

and

inapertus

Endoptychum depressum

], while

aridicola

Gyrophragmium dunalii

A. wariatodes

and

A. chartaceus

have affinities to taxa in sect.

Minores

(Mitchell and Bresinksy 1999, Geml 2004,

Gube 2009). The majority of taxa in these two groups

can be characterized by the yellow color change and

almond-like fragrance of the sporocarp (Moncalvo

et al. 2000, Geml 2004, Geml et al. 2004). Section

Arvenses

is known to be dominant in high-latitude

habitats, seemingly well adapted to harsh climates,

such as the subantarctic islands of New Zealand and

now the drier regions of Australia (Geml et al. 2007,

2008).

The position of

Agaricus melanosporus

and

Agaricus

inilleasper

are unresolved in DNA analyses (SUPPLE-

MENTARY FIG

.1,FIG. 1), and morphological characters

are not sufficiently distinct to place these taxa within

any particular section. Section

Sanguinolenti

includes

the sequestrate

A. erythrosarx

and agaricoid

A. lilaceps

A. pattersonae

A. fuscovelatus

and an undescribed

Australian species ‘‘C’’. All taxa in this clade have a

brownish pileus (

A. erythrosarx

is pale), rufescent

context and a fruity or spicy odor (Geml et al. 2004).

Thiers (1984) suggested that gastroid forms evolved

from an agaricoid ancestor via secotioid intermedi-

ates, considering the process to be sequential (i.e.

Cortinarius

Thaxterogaster

Hymenoga ster

)al-

though sometimes incomplete (i.e.

Agaricus

Endoptychum

). Under that hypothesis none of the

series examined here, in three different lineages

(

Macrolepiota

Chlorophyllum

Agaricus

), are complete

to the gastroid form, ending instead with the

secotioid taxa. However the more gastroid forms of

Barcheria

and

Endoptychum arizonicum

are unique

with no apparent extant secotioid or agaricoid sister

taxa. This may be due to extinction of related

sporocarp forms/taxa or reflect our incomplete

knowledge of local fungal diversity.

Timing of sequestration.—

Calibrating fungal phyloge-

nies is difficult due to lack of well preserved fossil

material (Matheny et al. 2009), so we incorporated an

existing hypothesis for a divergence time between

known species of

Agaricus

and

Chlorophyllum

from

Geml et al. (2004), which is derived from Heckman

et al. (2001). Divergence times of other taxa then are

calibrated based on this node, and our conclusions

about patterns of evolution of sequestrate forms

therefore are tentative at this stage. It should be

noted in particular that conclusions of Heckman et al.

(2001) may be overestimates of node ages (Nagy et al.

2011), such that our estimates here are possibly

conservative with respect to the dated origin of

sequestrate forms. New information about other

divergence times and/or fossil taxa will improve the

accuracy and interpretation of future calibrations.

An unexpected result from the calibrated phylog-

eny (F

IG. 3) is the relatively old sequestrate

Barcheria

clade, , 65 Ma, and

Agaricus melansporous

, , 34 Ma.

One possibility is that their ancestors also were

sequestrate, although this was not indicated by the

ancestral state reconstruction (F

IG.3,SUPPLEMENTARY

FIG

. 2). This then would have suggested that most

Agaricus

species have transitioned from sequestrate to

agaricoid forms, which is unlikely, but the evolution

of the gastroid form could be reversible (Hibbet

2004). More likely is that the lack of agaricoid forms

on the branches to

Barcheria

and

A. melanosporous

is a

result of incomplete taxon sampling and/or extinc-

tion of non-sequestrate forms.

Gastroid forms may have higher net rates of

diversification (Wilson et al. 2011), and their relative

taxonomic rarity may stem from recent origins.

Adaptation to xeric environments is often suggested

as the major selection pressure for fungal sporocarps

EBEL AND SYME:SEQUESTRATE

GARICUS

AND

ACROLEPIOTA

517

with an enclosed hymenium and eventually loss of the

stipe, and our calibrated phylogeny (F

IG. 3) suggests

that climatic events in Australia could be correlated

with the evolution of sequestrate forms. For taxa in this

study the evolution of sequestrate forms is estimated to

have occurred predominantly during the mid-Miocene

onward, during the past 15 000 000 y. Interestingly, a

recently calibrated phylogeny that includes one Aus-

tralian sequestrate form (

Auritella geoaustralis

) dates

the origin of its branch at a similar time frame, 20 Ma

(Matheny et al. 2009). This time frame matches

environmental and geological events, particularly for

Australia where the continent was subject to increased

cooling and drying as it moved northward (Martin

2006). Species radiations also occurred in the Miocene

for many important Australian sclerophyll flora, such

Eucalyptus

Banksia

and

Allocasuarina

(Crisp et al.

2004). Further study of Agaricaceae from xeric habitats

in Australia will be relevant for research into adaptive

evolutionary changes, particularly those that might be

driven by climatic shifts.

CKNOWLEDGMENTS

We thank the staff of PERTH, BRI, MEL and BPI for

providing access to specimens and aid in locating

collection data. Thank-you to Christopher Dunk (research

assistant) who aided with DNA extraction and sequencing,

and Tom May and the RBG Melbourne fungal studies

group for comment on the draft manuscript. Part of this

research was supported by the Australasian Biological

Resources Study research grant program. Finally many

thanks to reviewers who provided insightful commentary

and for suggesting a variation of a new taxon name

(erythrosarx).

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520 MYCOLOGIA

Agaricus eburneocanus voucher H7528 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2307 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher P32 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus erythrosarx voucher H7638 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher Trappe25186 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL1644 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher Watling14576 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus melanosporus voucher AD9763 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2155 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus melanosporus voucher AD9767 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher Trappe28630 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2308 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus erythrosarx voucher MURU6080 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher TWM1589 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Chlorophyllum agaricoides voucher Ginns11170 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher MEL2177563 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2057a internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2306 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2087 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus inilleasper voucher H4452 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Chlorophyllum agaricoides voucher Ginns11254 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher MEL2177561 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher H7518 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher H7564 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus erythrosarx voucher H7638 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2321 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus melanosporus voucher AD9763 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher P59 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher PMCatch540 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher H0219 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus inilleasper voucher H4452 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2154 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2058 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus chartaceus voucher H6271 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher DunstanL055 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota vinaceofibrillosa voucher H0047 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher TWM1589 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher MEL2058664 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher H7564 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher P36 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher P87 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher MEL2063323 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher MEL1053064 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota gasteroidea voucher H0052 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus eburneocanus voucher MURU5927 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Chlorophyllum agaricoides voucher AD3140 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus chartaceus voucher PERTH05848261 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher P71 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher DunstanL055 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher H7518 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL1650 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Macrolepiota turbinata voucher H0219 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2310 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher MEL260703 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2305 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus erythrosarx voucher MURU6080 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2513 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Barcheria willisiana voucher MEL269353 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus eburneocanus voucher H7528 28S ribosomal RNA gene, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL3005 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus wariatodes voucher PMCatch540 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus sp. TL2086 internal transcribed spacer 1, partial sequence; 5.8S ribosomal RNA gene, complete sequence; and internal transcribed spacer 2, partial sequence

Nucleotide Sequence

March 2011

Teresa Lebel · Anna E Syme

Agaricus radiatosquamulosus sp. nov. of A. subg. Spissicaules from Pakistan

Article

Jan 2023

In recent studies, new species and sections were described in Agaricus subg. Spissicaules. We intended to update this subgenus by including all these recent taxa, some interesting sequences retrieved in GenBank, and two new collections from the Punjab Province, Pakistan. Both morphological and phylogenetic approaches were used. Maximum Likelihood tree based on rDNA ITS sequences revealed that the sequences of the new specimens from Pakistan form a clade with five sequences from India retrieved from GenBank. For the seven samples of this clade, we described the new species Agaricus radiatosquamulosus based on the morphology of the two collections from Pakistan. This new species, which belongs to the recently proposed A. sect. Fulventes was compared to the three other species of this section. Except for the small-sized A. nanofulvus, the other species are difficult to recognize without DNA sequence data. The four species differ from each other at a poly T in ITS2. In addition, each of them is distributed on different continents. Agaricus radiatosquamulosus is the first report of a species of A. subg. Spissicaules in Pakistan. In addition, a sequence from GenBank identified as A. lanatoniger matches with another sample from New Zealand in A. sect. subrutilescentes. In absence of formally published taxonomic treatment, we provisionally named these samples A. cf. lanatoniger.

Phylogeny of Agaricus subgenus Pseudochitonia with the description of a new section and a new species from Oman

Article

Full-text available

Aug 2022

Agaricus is an important mushroom genus with some most common edible and medicinal species, but some members are also poisonous. Taxonomically, the genus is divided into seven subgenera including the subgenus Pseudochitonia. In this study,we analyzed some collections of Agaricus from southern Oman, belonging to A. subgenus Pseudochitonia. Agaricus subg. Pseudochitonia is the largest subgenus in terms of sections, with 13 sections by applying the modern taxonomic system in which supraspecific taxa are ranked according to their divergence times. We applied the same procedure using BEAST (Bayesian evolutionary analysis sampling tree) to evaluate the taxonomic position of our specimens. Our specimens formed a strong monophyletic clade with unclassified Agaricus species (A. lannaensis and A. sp. CA820), based on multigene sequence data (ITS, 28S and TEF1) and ITS alignment. The divergence time is estimated around 20–30 million years for this clade. We proposed a new section Catenulati following the basic principles of modern taxonomic system of Agaricus, which are (i) strong phylogenetic support, (ii) divergence times younger than 30 Ma, and (iii) specific morphological characters where possible. The new section Catenulati is represented by a new species Agaricus arabiensis. Moreover, the new section resolved the pending issue of A. lannaensis, previously as unclassified taxon in A. subg. Pseudochitonia and is now in A. sect. Catenulati. The new section is characterized by a phenolic odor of basidiomata and catenoid cheilocystidia.

Agaricus subgenus Pseudochitonia in Malakand, Pakistan: An updated phylogeny and description of three new species

Article

Apr 2024
MYCOLOGIA

Agaricus is a genus with more than 500 species. Most of the new species reported since 2000 are tropical or subtropical. The study area, the Malakand region, located in the north of Pakistan, has a subtropical climate. In this study, nine species, including three new species, of Agaricus subgenus Pseudochitonia, are reported from this region. Description of the new species are based on morphological characteristics and phylogenetic analyses using three DNA regions: nuc ribosomal DNA internal transcribed spacers (ITS), fragments of the large subunit of nuc ribosomal DNA (28S), and the translation elongation factor 1-alpha gene (TEF1). One new species, Agaricus lanosus, with wooly squamules on its cap, forms a lineage within Agaricus sect. Bivelares and cannot be classified with certainty in one of the two subsections (Cupressorum and Hortenses) of this section. Agaricus rhizoideus with rhizoid-like structure at the base of the stipe forms a basal clade in Agaricus sect. Hondenses. Specimens of the third new species, Agaricus malakandensis, form a species-level clade within Agaricus sect. Catenulati and exhibits the morphological characteristics of this section. Due to their similar ITS sequences, two previously unnamed specimens from Thailand (A. sp. LD2012162 and CA799) are considered conspecific with A. malakandensis.

Agaricus cervinoculus sp. nov. (Agaricaceae), a new wild edible mushroom from Mexico

Article

Full-text available

Apr 2024

In Mexico, there are 55 wild species of Agaricus representing approximately 10 % of the world’s diversity of the genus. 22 species of these wild mushrooms are traditionally consumed. The lack of keys for the identification of Mexican species, the difficulties in recognizing distinctive morphological characters and the need for comprehensive studies in some sections of the genus, makes it difficult to distinguish species complexes. The edible species known locally as “ojo de venado” was previously identified as A. sylvaticus in Mexico. Our phylogenetic analyses revealed that this ethnotaxon, which is sold in the markets of Xalapa (Veracruz, Mexico), includes at least two species of A. sect. Subrutilescentes. One of them, Agaricus cervinoculus, is described as a new species. A full description, photographs and comparisons to other similar species are provided. This new species constitutes a well-supported monophyletic clade, sister to the “A. subrutilescens” group.

Fungal diversity notes 1717–1817: taxonomic and phylogenetic contributions on genera and species of fungal taxa

Article

Full-text available

Feb 2024
FUNGAL DIVERS

As the continuation of Fungal Diversity Notes series, the current paper is the 16th contribution to this series. A total of 103 taxa from seven classes in Ascomycota and Basidiomycota are included here. Of these 101 taxa, four new genera, 89 new species, one new combination, one new name and six new records are described in detail along with information of hosts and geographic distributions. The four genera newly introduced are Ascoglobospora, Atheliella, Rufoboletus and Tenuimyces. Newly described species are Akanthomyces xixiuensis, Agaricus agharkarii, A. albostipitatus, Amphisphaeria guttulata, Ascoglobospora marina, Astrothelium peudostraminicolor, Athelia naviculispora, Atheliella conifericola, Athelopsis subglaucina, Aureoboletus minimus, A. nanlingensis, Autophagomyces incertus, Beltrania liliiferae, Beltraniella jiangxiensis, Botryobasidium coniferarum, Calocybella sribuabanensis, Calonarius caesiofulvus, C. nobilis, C. pacificus, C. pulcher, C. subcorrosus, Cortinarius flaureifolius, C. floridaensis, C. subiodes, Crustomyces juniperi, C. scytinostromoides, Cystostereum subsirmaurense, Dimorphomyces seemanii, Fulvoderma microporum, Ginnsia laricicola, Gomphus zamorinorum, Halobyssothecium sichuanense, Hemileccinum duriusculum, Henningsomyces hengduanensis, Hygronarius californicus, Kneiffiella pseudoabdita, K. pseudoalutacea, Laboulbenia bifida, L. tschirnhausii, L. tuberculata, Lambertella dipterocarpacearum, Laxitextum subrubrum, Lyomyces austro-occidentalis, L. crystallina, L. guttulatus, L. niveus, L. tasmanicus, Marasmius centrocinnamomeus, M. ferrugineodiscus, Megasporoporia tamilnaduensis, Meruliopsis crystallina, Metuloidea imbricata, Moniliophthora atlantica, Mystinarius ochrobrunneus, Neomycoleptodiscus alishanense, Nigrograna kunmingensis, Paracremonium aquaticum, Parahelicomyces dictyosporus, Peniophorella sidera, P. subreticulata, Phlegmacium fennicum, P. pallidocaeruleum, Pholiota betulicola, P. subcaespitosa, Pleurotheciella hyalospora, Pleurothecium aseptatum, Resupinatus porrigens, Russula chlorina, R. chrysea, R. cruenta, R. haematina, R. luteocarpa, R. sanguinolenta, Synnemellisia punensis, Tenuimyces bambusicola, Thaxterogaster americanoporphyropus, T. obscurovibratilis, Thermoascus endophyticus, Trechispora alba, T. perminispora, T. subfarinacea, T. tuberculata, Tremella sairandhriana, Tropicoporus natarajaniae, T. subramaniae, Usnea kriegeriana, Wolfiporiella macrospora and Xylodon muchuanensis. Rufoboletus hainanensis is newly transferred from Butyriboletus, while a new name Russula albocarpa is proposed for Russula leucocarpa G.J. Li & Chun Y. Deng an illegitimate later homonym of Russula leucocarpa (T. Lebel) T. Lebel. The new geographic distribution regions are recorded for Agaricus bambusetorum, Bipolaris heliconiae, Crinipellis trichialis, Leucocoprinus cretaceus, Halobyssothecium cangshanense and Parasola setulosa. Corresponding to morphological characters, phylogenetic evidence is also utilized to place the above-mentioned taxa in appropriate taxonomic positions. The current morphological and phylogenetic data is helpful for further clarification of species diversity and exploration of evolutionary relationships in the related fungal groups.

Agaricus section Minores: New and noteworthy species from India

Article

Jan 2024

In this study, four species of Agaricus sect. Minores are presented based on morphological and nrITS based phylogenetic analyses (Maximum Likelihood and Bayesian Inference). The molecular analysis and morphological characteristics supported the placement of these species in A. sect. Minores. Three species, viz., Agaricus indicus, A. palodensis, and A. waya-nadensis are new to science, while A. purpureofibrillosus is a new phytogeographical record. Comprehensive descriptions, photographs, phylogenetic analyses results, and comparisons with related species are provided.

Paedomorphosis and Evolution of Sequestrate Basidiomycetes

Chapter

Full-text available

Aug 2023

Current theories on the evolution of sequestrate (enclosed) basidiomes explain the origin of these forms in a gradualist adaptational process led by selective forces, such as drought and animal consumption. Paedomorphosis (the retention of juvenile traits) has been invoked as the phenomenon underlying sequestration, but many consequences of this process have not yet been explored. Our present interpretation of sequestrate morphologies, in light of Stephen Jay Gould’s characterization of neoteny (retention of juvenile features in an adult stage with mature reproductive structures) and progenesis (the onset of sexual maturity in a morphologically immature stage that does not reach the mature morphology observed in the ancestral form), both involved in paedomorphosis, implies that the origin of sequestrate basidiomes might constitute two distinct evolutionary processes. These two processes could be recognized among fungi by contrasting their morphological plasticity, phylogenetic diversification, and ecological patterns. The hypotheses discussed here provide new insights for interpreting and studying the evolution of sequestrate fungi.KeywordsGasteroidSecotioidProgenesisNeotenyHeterochrony

A new species of Agaricus sect. Agaricus (Agaricaceae) from India

Article

Full-text available

Nov 2022
NORD J BOT

Agaricus albovariabilis is presented as a new species from peninsular India on the basis of morphological features and ITS sequence analysis. It is charcterized by small to medium sized white basidiomata, becoming brownish at maturity, unchanging context, mild anise odor, negative Schäffer’s and KOH reactions, and clavate to vesiculose clavate cheilocystidia. It is presented with detailed descriptions, photographs, phylogenetic reconstruction and a comparison with phenetically similar species.

Chlorophyllum molybdites, specie tossica d'oltre oceano ambientata in Sicilia

Article

Full-text available

Oct 2022

Angelo Miceli

Introduzione In questo contributo viene trattato un taxon esotico, tipico e molto diffuso nei paesi extraeuropei quali la parte meridionale del Nord America, l'America centrale, l'America del sud, l'Africa, il Madagascar, l'Oceania fino alle isole del Pacifico [Brunelli, 2009]. Alla luce dei numerosi ritrovamenti degli ultimi anni, possiamo affermare che questa specie sia solita fruttificare in maniera abbondante e con una certa frequenza nella zona costiera della Sicilia orientale, un fenomeno che nella nostra regione si può ad oggi riscontrare per numerose altre specie ad analoga tipicità. Chlorophyllum molybdites ha una conformazione morfologico-strutturale molto simile a tutte le Macrolepiota del gruppo rhacodes, oggi inserite nel genere Chlorophyllum, ma in particolare la somiglianza è condivisa con la più nota Macrolepiota procera, buon commestibile, con la quale viene facilmente confusa dai cercatori meno esperti. La confusione può essere pericolosa in quanto C. molybdites è specie tossica che causa un tipo di sindrome definita Morgana (1). Il suo primo ritrovamento per l'Europa, avvenuto in ambiente naturale e ampiamente documentato, risale all'anno 2005 e fa riferimento a sporofori ritrovati in Sicilia, nel territorio del comune di Mascali, in provincia di Catania [Signorino, 2006a, 2006b; Signorino, 2015a, 2015b]. La prima intossicazione dovuta al suo consumo viene registrata nell'anno 2014 e coinvolge due persone (madre e figlio), ricoverate per le cure del caso presso l'Ospedale di Giarre (CT) [Signorino, 2015a, 2015b]. Genere Chlorophyllum Massee Bull. Misc. Inf. Kew (n. 138): 135 (1898) Il genere, la cui specie tipo è Chlorophyllum molybdites (G. Mey.) Massee, ospita sporofori più o meno carnosi, di taglia medio-grande [La Chiusa, 2013], a portamento lepiotoide che, come già detto, nell'aspetto morfo-cromatico generale manifestano notevoli similarità con le diverse specie del genere Macrolepiota Singer, in particolare con quelle appartenenti al gruppo rhacodes. I rappresentanti di questo genere presentano le seguenti caratteristiche: 1) Sindrome Morgana, conosciuta anche come Morganismo, è una sindrome a breve latenza che causa disturbi gastrointestinali di notevole entità. I sintomi dell'intossicazione si presentano tra 30 minuti e 2-4 ore dal consumo [Assisi et al., 2008; Della Maggiora & Pera, 2021], si manifestano con disturbi intensi e persistenti quali, ad esempio: pesantezza gastrica, nausea, conati di vomito ripetuti e violenti spesso accompagnati da emissione di sangue, coliche addominali con violente scariche diarroiche spesso sanguinolente, difficoltà respiratoria, vertigini, sudorazioni, riduzione delle capacità di ragionamento, crampi muscolari, colorito subitterico della cute, tachicardia, ipercromia delle urine, ipersecrezione lacrimare e salivale [Brunelli, 2009]. A volte si manifesta anche cianosi labiale e linguale. Il recupero è totale anche senza l'ausilio di una terapia specifica [Della Maggiora & Pera, 2021]. La denominazione della sindrome Morganismo, trova origine in uno dei sinonimi della specie responsabile dell'intossicazione: Lepiota morganii [Assisi et al., 2008].

Mua et al-Agaricus iesu et marthae

Article

Sep 2022

Riassunto Viene descritto ed illustrato Agaricus iesu-et-marthae, raccolto in Sardegna, che costituisce la prima segnalazione in Italia. Viene inoltre proposto un albero filogenetico relativo alle specie di Agaricus sez. Minores, clado I, sec. He et al. (2017). Abstract Agaricus iesu-et-marthae, found in Sardinia, which constitutes the first report in Italy, is described and illustrated. Is also proposed a phylogenetic tree relating to the species of Agaricus sect. Minores clade I, according to He et al. (2017). Introduzione Negli ultimi anni il genere Agaricus si è arricchito di un notevole numero di specie ed oggi se ne contano oltre 580 a livello mondiale. Precedentemente la distinzione tra le specie di questo genere era affidata esclusivamente ai caratteri macro e micromorfologici oltre che chimici e molto spesso era difficile separare specie molto simili tra loro (specie criptiche). La separazione tra queste specie è potuta avvenire grazie alla biologia molecolare; tale disciplina ha dato un contributo decisivo alla classificazione del genere Agaricus, apportando delle sostanziali modifiche, soprattutto grazie ai lavori di vari studiosi europei ed extraeuropei (Zhao et al. 2011, Zhao et al. 2016, Kerrigan 2016, Chen et al. 2017, Parra et al. 2018, He et al. 2018) che attualmente prevedono all'interno del genere 6 sottogeneri e 26 sezioni. Le specie appartenenti ad A. sez. Minores a livello mondiale sono da 85 a 100, numero che può variare considerando le sinonimie, ma si stima che il numero totale possa arrivare a 200 (Chen et al. 2017). Infatti soltanto dal 2016 a oggi sono stati descritti ben 43 taxa di A. sez. Minores: Li et al. 2016, con 2 specie, Parra et al.

Dictionary of the Fungi

Article

Full-text available

Jan 2009

The Lycoperdales. A molecular approach to the systematics of some gasteroid mushrooms

Article

Sep 2001

Based on selected taxa of Lycoperdales and Geastrales, phylogeny and delimitation of several gasteroid mushroom genera were investigated using RFLP and sequence analysis of 18S rDNA and ITS rDNA (ITS1-5.8S-ITS2). Support for an Agaricales-Lycoperdales relationship was found, the Geastrales being clearly separated from this lineage. Within the Lycoperdales, Lycoperdon was found to be polyphyletic, whereas Bovista appeared to be monophyletic.

Phylogenetic relationships of Agaricus species based on ITS-2 and 28S ribosomal DNA sequences

Article

Sep 1999

Parsimony and maximum likelihood analyses of sequence data from the internal transcribed spacer (ITS-2) region plus a portion of the 28S gene are used to study the phylogenetic relationships of Agaricus species. The ingroup contained 16 Agaricus species, selected from the range of sections contained in the subgenus Agaricus. A total of eight out-groups were chosen from the Agaricaceae and phylogenetic trees rooted on Stropharia coronilla. Results suggest that Agaricus may be monophyletic, but further tests should include taxa from the subgenera Langagaricus and Conioagaricus. Two well-supported clades were found. The Agaricus clade contained A. subperonatus, A. devoniensis, A. bisporus, A. spissicaulis, A. bitorquis, and A. impudicus. Results suggest that A. bernardii may be closely related to this group. The Arvenses clade contained A. silvicola, A. arvensis, A. abruptibulbus, and A. semotus. A possible sister group to the Arvenses clade includes A. lanipes and A. maskae. Relationships for A. campestris, A. silvaticus and A. xanthoderma remain unclear, although it is likely these species diverged early compared with the other Agaricus species studied. Phylogenetic hypotheses are given for taxa that have been difficult to classify based on other forms of data, including the placement of A. spissicaulis with A. bisporus; A. lanipes with A. maskae; and Melanophyllum haematospermum with Leucoagaricus naucinus.

The Secotioid Syndrome

Article

Jan 1984

Harry D. Thiers

The Agaricales in Modern Taxonomy.

Article

Jul 1987

Agaricus section Xanthodermatei : a phylogenetic reconstruction with commentary on taxa

Article

Nov 2005

Agaricus section Xanthodermatei comprises a group of species allied to A. xanthodermus and generally characterized by basidiomata having phenolic odors, transiently yellowing discolorations in some parts of the basidiome, Schaeffer’s reaction negative, and mild to substantial toxicity. The section has a global distribution, while most included species have distributions restricted to regions of single continents. Using specimens and cultures from Europe, North America, and Hawaii, we analyzed DNA sequences from the ITS1+2 region of the nuclear rDNA to identify and characterize phylogenetically distinct entities and to construct a hypothesis of relationships, both among members of the section and with representative taxa from other sections of the genus. 61 sequences from affiliated taxa, plus 20 from six (or seven) other sections of Agaricus, and one Micropsalliota sequence, were evaluated under distance, maximum parsimony and maximum likelihood methods. We recognized 21 discrete entities in Xanthodermatei, including 14 established species and 7 new ones, three of which are described elsewhere. Four species from California, New Mexico, and France deserve further study before they are described. Type studies of American taxa are particularly emphasized, and a lectotype is designated for A. californicus. Section Xanthodermatei formed a single clade in most analyses, indicating that the traditional sectional characters noted above are good unifying characters that appear to have arisen only once within Agaricus. Deep divisions within the sequence-derived structure of the section could be interpreted as subsections in Xanthodermatei; however, various considerations led us to refrain from proposing new supraspecific taxa. The nearest neighbors of section Xanthodermatei are putatively in section Duploannulati.

Phylogeny and taxonomy of Macrolepiota (Agaricaceae)

Article

May 2003

The position and composition of Macrolepiota within the Agaricaceae and its phylogenetic relationships with other members of the family were investigated, using both molecular (ITS and LSU rDNA sequences) and morphological characters. The molecular data separate the genus into two clades. The first clade comprises M. procera, M. mastoidea, M. clelandii and allies and is a sister group of Leucoagaricus and Leucocoprinus. The second, more diverse, clade, with M. rachodes and allies, M. globosa, Chlorophyllum molybdites, Leucoagaricus hortensis and Endoptychum agaricoides, is a sister group of Agaricus. The separation of the two clades is supported by morphological characters, such as the structure of the pileus covering, the stipitipellis and the shape of the germ pore and the spore apex. The two clades are regarded as genera for which the names Macrolepiota and Chlorophyllum are proposed. Macrolepiota nympharum does not belong to either clade but is assigned to the genus Leucoagaricus, close to L. leucothites. Endoptychum depressum is transferred to the genus Agaricus as A. inapertus.

New Australian fungi

Article

M.C. Cooke

A Catalogue of Leonard Rodway’s Collection of Fungi. 1. Hymenomycetes and Gasteromycetes

Article

Jan 1993

Calibrating molecular estimates of substitution rates and divergence times in birds

Article

Jul 2007

Simon Y. W. Ho

Many evolutionary studies of birds rely on the estimation of molecular divergence times and substitution rates. In order to perform such analyses, it is necessary to incorporate some form of calibration information: a known substitution rate, radiometric ages of heterochronous sequences, or inferred ages of lineage splitting events. All three of these techniques have been employed in avian molecular studies, but their usage has not been entirely satisfactory. For example, the 'traditional' avian mitochondrial substitution rate of 2% per million years is frequently adopted without acknowledgement of the associated uncertainty. Similarly, fossil and biogeographic information is almost always converted into an errorless calibration point. In both cases, the resulting estimates of divergence times and substitution rates will be artificially precise, which has a considerable impact on hypothesis testing. In addition, using such a simplistic approach to calibration discards much of the information offered by the fossil record. A number of more sophisticated calibration methods have recently been introduced, culminating in the development of probability distribution-based calibrations. In this article, I discuss the use of this new class of methods and offer guidelines for choosing a calibration technique.

Sequestrate species of Agaricus and Macrolepiota from Australia: New species and combinations and their position in a calibrated phylogeny

Abstract and Figures

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