Geastrum Sect Geastrum 1422974412556

Persoonia 34, 2015: 130 –165
www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE http://dx.doi.org/10.3767/003158515X687443
Integrative taxonomy reveals an unexpected diversity in

Geastrum section Geastrum (Geastrales, Basidiomycota)
J.C. Zamora 1*, F.D. Calonge 1, M.P. Martín 1
Key words Abstract   A revision of the classification of Geastrum sect. Geastrum is presented on the basis of an integrative
taxonomic approach, which involves the study of morphological, molecular, ecological, and chorological data. Four
chorology
DNA regions are analysed: the ITS and LSU nrDNA, rpb1, and atp6. Phylogenetic reconstructions include 95 ingroup
ecology
samples and show five main clades, which are considered as five subsections, one of them proposed as new (G. sub-
Geastraceae
sect. Hungarica), and a total of 27 lineages recognizable at species level. Discriminant function analyses, ANOVAs
morphology
and Tukey’s HSD tests on more than 500 basidiomata reveal the utility of several quantitative morphological char-
phylogenetics
acters for species delimitation. As a result of the combination of the different sources of taxonomic information, a
species delimitation
revised taxonomy is presented and seven new species, viz., G. austrominimum, G. benitoi, G. britannicum, G. ku-
harii, G. meridionale, G. papinuttii, and G. thanatophilum, are proposed.
Article info   Received: 13 June 2014; Accepted: 20 October 2014; Published: 3 February 2015.
INTRODUCTION It is known that different species delimitation approaches can

produce highly deviant results, thus producing taxonomic con-
Taxonomy of earthstars (genus Geastrum) has been tradition- flicts that destabilise systematics. What is more, all methods
ally based on morphological traits of the basidiomata. Among have weaknesses and may fail in species delimitation (Schlick-
the many studies about the taxonomy of the genus Geastrum, Steiner et al. 2010, Carstens et al. 2013). Instead of fighting
Sunhede’s monograph (Sunhede 1989) is by far the most com- between two of the most used methods for species delimita-
prehensive one. His taxonomic view and species circumscription, i.e., phylogenetic and morphological species recognition,
tion are followed by most recent authors, sometimes with minor the combination of data from these approaches may result in
exceptions (e.g., Calonge 1998, Sarasini 2005, Jeppson 2013). more stable classifications accessible for a larger number of
However, a more complex scenario is coming to light with the researchers. The sum of several taxonomic information sources
inclusion of molecular phylogenetic data. For instance, Kasuya has been sometimes called ‘integrative taxonomy’, a term firstly
et al. (2012) found an unexpected phylogenetic diversity hid- proposed by Dayrat (2005). However, Dayrat’s concept is rather
den under an a priori ‘single morphological species’ (G. triplex strict and some of his guidelines are criticable, as Valdecasas
s.l.). The systematics of the whole genus Geastrum concerning et al. (2008) carefully explained. Even if it is not always easy to
infrageneric subdivisions is being elucidated thanks to DNA- find balance among the different data, the results may reflect
based molecular data (Zamora et al. 2014), and the necessity a more natural and accurate classification of the involved taxa
of finding new morphological or chemical information sources (Ruiz-Sánchez & Sosa 2010, Medina et al. 2012, Edwards &
to determine the phylogenetic and taxonomic boundaries has Knowles 2014).
been also pointed out (Zamora et al. 2013, 2014). Among the 14 sections of Geastrum considered in Zamora et al.
Morphologically close taxa are an important source of taxo- (2014), we decided to focus the present study on G. sect.
nomic problems and disagreement in classificatory systems. Geastrum, which includes species with a glabrous endo
So, it is necessary that adequate samples of specimens are peridial surface, coated by a mesoperidial layer made up by
available, in order to properly identify reliable differences and generative hyphae and crystalline matter; slender, ellipsoid
not only the extremes of the intraspecific variation. In these to cylindrical basidia; double-layered mycelial layer encrust-
cases, multivariate analysis based on large sets of morphologi- ing debris; basidiospores with verrucose to baculate-pilate
cal data have proven to be useful for taxonomic purposes, by ornamentation; and rhizomorphs with mostly calcium oxalate
identifying morphologically homogeneous groups (Pimentel dihydrate (COD) bipyramidal crystals, sometimes mixed with
1979, Valcárcel & Vargas 2010, Jiménez-Mejías et al. 2014). calcium oxalate monohydrate (COM) horn-like ones (G. ovali
In particular, linear discriminant function analyses have been sporum, G. parvistriatum, G. pectinatum, G. striatum), but rare-
used in Botany for decades to identify useful characters that ly only horn-like (G. coronatum) (Zamora et al. 2013). This
allow the distinction of different taxa (Henderson 2006), being section was subdivided into four subsections, distinguishable
the ‘Fisher’s Iris dataset’ the classical example of this type of by some macromorphological traits (mainly the peristome
analysis (Fisher 1936). However, few studies in Mycology have type and the mesoperidium structure) as well as features of
performed discriminant analyses for assessing morphological the basidiospores. Finally, G. hungaricum was indicated by
differences among species (Smith et al. 2004, Roca-Valiente Zamora et al. (2014) to possibly belong to this section, based
2013), despite they are very promising for morphology-based on morphology and molecular data of Jeppson et al. (2013),
species identification. but it was not definitely included awaiting further data.
1
Real Jardín Botánico-CSIC, Plaza de Murillo 2, E-28014, Madrid, Spain. Some of these species are considered to be well-known and ap-
*
Corresponding author e-mail: jcsenoret@gmail.com. parently widespread (e.g., G. coronatum, G. minimum, G. pecti-
© 2015 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures

You are free to share - to copy, distribute and transmit the work, under the following conditions:
Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work).
Non-commercial: You may not use this work for commercial purposes.
No derivative works: You may not alter, transform, or build upon this work.
For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be
waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights.
J.C. Zamora et al.: Diversity in Geastrum section Geastrum 131
natum, G. quadrifidum, G. striatum), while some others are still s.l., G. glaucescens s.l., G. leptospermum, G. minimum s.l., G.
undescribed. Paying special care to the literature, numerous ovalisporum, G. parvistriatum, G. pectinatum s.l., G. quadri-
taxonomic problems arise around different species included fidum, G. smithii s.l., and G. striatum. Geastrum hungaricum
in G. sect. Geastrum. The so-called G. minimum is normally was also added to test whether it nests or not in this section,
treated as a single, rather variable species, mostly small-sized since previous phylogenies have shown this species more or
and often showing quite big crystals of calcium oxalate on the less closely related to some G. sect. Geastrum species (Jepp-
endoperidial surface. Under this broad concept many names son et al. 2013), and also some morphological traits suggested
have been included as synonyms, such as the South African it belonging to sect. Geastrum (Zamora et al. 2014). Finally,
G. calceum, the North American G. juniperinum, and the Euro- additional herbarium specimens not included in Zamora et al.
pean G. marginatum, G. cesatii, G. granulosum, and G. mini- (2014), but with morphological characters that match to those
mum var. fumosicollum (Cunningham 1944, Bottomley 1948, that define the sect. Geastrum, were added. The studied
Sunhede 1989). Zamora et al. (2014) showed at least two specimens came from the following herbaria: AH, B, BAFC,
well-separated phylogenetic lineages under the name ‘G. mini- BPI, CANB, CORD, CUP, K, LPS, MA-Fungi, MEL, MICH,
mum agg.’, as well as samples provisionally determined as PRM, S, UFRN, UPS, and the private herbaria of M.Á. Ribes, S.
G. calceum. Sunhede, J.C. Zamora, Asociación Vallisoletana de Micología
Geastrum pectinatum is another widespread taxon that may (AVM), and Sociedad Micológica Errotari (ERRO).
involve more than one species. Palmer (1959) studied the type Ninety five ingroup specimens were included in our molecular
specimens of G. plicatum and G. tenuipes, concluding that analyses (Table 1). A special effort was made in sampling spe-
both can be included under the variation of G. pectinatum, and cimens of G. minimum s.l. and G. pectinatum s.l., since Zamora
Sunhede (1989), based on that study and on the protologues of et al. (2014) revealed various well-defined phylogenetic line-
these species, agreed with the synonymy. Geastrum biplicatum ages for each of these taxa. Two specimens of G. fornicatum
and G. calyculatum are also often considered synonyms (Cun- were used as outgroup based on Zamora et al. (2014). To mini-
ningham 1944, Bottomley 1948). Zamora et al. (2014) showed mize the inclusion of missing data, molecular data of specimens
a single clade containing samples from various worldwide present in the GenBank database not generated by us were
locations, but with a considerable variation in sequence data. only included for certain groups (especially the G. minimum
The South European and recently proposed G. parvistriatum group and the G. pectinatum group), when either more than
(Zamora & Calonge 2007) has been carefully compared with half of the DNA regions used were available for the same
other morphologically close European taxa, particularly to G. stri- specimen, or when we lack newly generated sequences of a
atum, and the differences of both species are clearly established particular species.
by both morphological and molecular phylogenetic data (Jepp- A total of 565 mature basidiomata were measured for morpho-
son et al. 2013; Zamora et al. 2014). However, Zamora et al. metric analyses, and additional specimens not included in these
(2014) included sequences of the Argentinian G. glaucescens, a analyses were examined to properly describe the proposed new
species that seems to be phylogenetically very close to G. par- species. Complete descriptions are provided in addition to the
vistriatum. Kuhar et al. (2013) provided a rather detailed de- diagnosis for all new species, and only synoptic descriptions
scription of G. glaucescens and pointed out the morphological with the most relevant or diagnostic characters that define each
similarities with G. parvistriatum, concluding that they are dif- taxon were included for the already described species. Those
ferent species despite their close relationship. synoptic descriptions are based on both the literature (then
There are other minor taxonomic problems in this group. For in- references are provided) and newly observed data. Terminology
stance, G. arenarium, originally described from North America, mostly followed Sunhede (1989).
has been considered a rather widespread taxon, having been Due to the high amount of revised herbarium collections, for
recorded from Australia (Cunningham 1944), South Africa most taxa we only cite specimens used for molecular or mor-
(Bottomley 1948), South Europe (Calonge & Zamora 2003), phological analyses. Specimens that significantly contributed
and South America (Kuhar et al. 2013). However, the icono to the intraspecific variation are also mentioned. In the case of
graphy and descriptions of the South European specimens are the new species, all examined specimens are cited.
somewhat deviating because of the exoperidial rays with a less Ecological data were taken from the literature, herbarium labels,
marked hygrometry (Calonge & Zamora 2003, Jeppson 2013). and our own observed data from newly collected specimens. Ter-
Finally, G. striatum is often treated as a very well-defined taxon minology of biomes and ecozones follows Olson et al. (2001).
due to its unique ring-like apophysis (Sunhede 1989), but some
extra-European records are based on specimens with not so Molecular analyses
well-developed apophyses (e.g., Ochoa & Moreno 2006), and
Methodology concerning molecular analyses followed Zamora
may represent different species.
et al. (2014) and therefore it is summarized next.
Thus, the main goal of this study is to clarify the systematics
The following DNA regions were studied: ITS (including ITS1,
of Geastrum sect. Geastrum using both molecular and mor-
5.8S, and ITS2) and 28S (LSU) nrDNA, rpb1, and atp6. Primers
phological approaches, also taking into account other sources
used for PCR amplification of the target fragments were: ITS1F
of information as well, such as chorology and ecology. This
(Gardes & Bruns 1993) and ITS4 (White et al. 1990) for the ITS
integrative approach lead us to describe the new taxa found
region; LR0R (Cubeta et al. 1991) and LR5 (Vilgalys & Hester
during this study, determining whether the different phylogenetic
1990) for LSU; gRPB1A (Stiller & Hall 1997)/fRPB1C (Matheny
clades obtained through molecular data can be distinguished
et al. 2002) and/or RPB1GEA-1F/RPB1GEA-2r (Zamora et al.
on account of their morphological characteristics.
2014) for rpb1; and atp6-1, atp6-2, atp6-3 (Kretzer & Bruns
1999), ATP6GEA-ir, and ATPGEA-iF (Zamora et al. 2014)
MATERIAL AND METHODS for atp6. Sometimes, ITS and LSU were amplified together,
and then the primers used were ITS1F and LR5. PCR cycling
Study group and sampling strategy parameters follow Martín & Winka (2000) for ITS, Zamora et
The study group comprises all species and species complexes al. (2014) for LSU (alone or together with ITS) and rpb1, and
included by Zamora et al. (2014) as members of Geastrum sect. Kretzer & Bruns (1999) for atp6. PCR products were purified
Geastrum, i.e., G. arenarium s.l., G. calceum s.l., G. coronatum with ExoSap-IT® (VWR, Spain) or using the QIAquick® Gel
132 Persoonia – Volume 34, 2015
Table 1   Summary of specimens included in molecular analyses, for which geographical origin, herbarium vouchers, and GenBank accession numbers for
each DNA region are provided. New sequences generated in this study are marked in bold.
Taxon Country and state /province Herbarium voucher GenBank accession numbers
ITS LSU rpb1 atp6
G. arenarium Argentina, La Rioja MA-Fungi 83760 KF988351 KF988471 KF988606 KF988741

G. austrominimum Australia, New South Wales CANB 748741 – KP687529 KP687531 KP687572
Australia, New South Wales MEL 2276089 KP687490 KP687451 KP687532 KP687573
Australia, Victoria MEL 2292062 KP687491 KP687452 KP687533 KP687574
G. benitoi Spain, Madrid MA-Fungi 68191 KF988350 KF988469 KF988604 KF988739
Spain, Madrid MA-Fungi 87324 KP687494 KP687455 KP687536 KP687577
Spain, Madrid Zamora 76 KF988338 KF988470 KF988605 KF988740
G. cf. biplicatum Japan, Mie TNS Sakamoto 182 JN845113 JN845231 – JN845355
Japan, Shizuoka TNS Sakamoto 191 JN845114 JN845232 – JN845356
Japan, Aomori TNS TKG-GE-91003 JN845110 JN845228 – JN845352
Japan, Shizuoka TNS TKG-GE-91102 JN845115 JN845233 – JN845357
G. britannicum England, Hampshire K(M) 60288 EU784242 – – –
England, Norfolk K(M) 79617 EU784243 – – –
England, Norfolk K(M) 99914 EU784244 – – –
G. cf. calceum1 Argentina, Tucumán MA-Fungi 83761 KF988341 KF988478 KF988613 –
G. cf. calceum2 Brazil, Rio Grande do Norte UFRN-Fungos 723 KF988340 KF988477 KF988612 KF988747
G. coronatum Hungary, Nagyerdö Csokás PRM 842868 KP687495 KP687456 KP687537 –
Spain, Madrid Zamora 181 KP687496 KP687457 KP687538 KP687578
Spain, Sweden Zamora 522 KF988362 KF988484 KF988619 KF988754
USA, Arizona MICH 28567 KF988363 KF988485 KF988620 KF988755
G. fornicatum Spain, Lérida MA-Fungi 30749 KF988375 KF988497 KF988632 KF988767
Spain, Valladolid Zamora 255 KF988374 KF988496 KF988631 KF988766
G. glaucescens Argentina, La Rioja MA-Fungi 83762 KF988378 KF988500 KF988635 KF988770
Argentina, Catamarca MA-Fungi 83763 KF988379 KF988501 KF988636 KF988771
G. granulosum Russia, Rostov K(M) 154623 JN845105 JN845223 – JN845347
USA, Arizona MICH 28119a KP687498 KP687459 KP687540 KP687580
USA, Arizona MICH 28210 KP687499 KP687460 KP687541 KP687581
USA, Wisconsin MICH 72010 KF988402 KF988530 KF988665 KF988797
Sweden, Öland Sunhede 7746 KF988401 KF988529 KF988664 KF988796
G. hungaricum Hungary GB MJ8915 KC581964 KC581964 – –
Slovakia GB MJ9317 KC581963 KC581963 – –
Czech Republic, Reporyje Sunhede 5993 KP687500 KP687461 KP687542 KP687582
Spain, Toledo Zamora 611 KP687501 KP687462 KP687543 KP687583
G. kuharii Argentina, Buenos Aires MA-Fungi 83795 KF988463 KF988598 KF988733 KF988864
Argentina, Entre Ríos MA-Fungi 86913 KP687502 KP687463 KP687544 KP687584
Argentina, Buenos Aires MA-Fungi 86914 KP687503 KP687464 KP687545 KP687585
G. marginatum Spain, Canary Islands ERRO 2012112609 KP687504 KP687465 KP687546 KP687586
Spain, Madrid MA-Fungi 31530 KF988404 KF988532 KF988667 KF988799
Spain, Jaén MA-Fungi 32395 KP687505 KP687466 KP687547 KP687587
Sweden, Gotland MA-Fungi 86669 KF988405 KF988533 KF988668 KF988800
USA, Arizona MICH 28119b KF988403 KF988531 KF988666 KF988798
Czech Republic, Bohemia PRM 842884 KP687507 KP687468 KP687549 –
G. meridionale Spain, Valladolid AVM 2708 KP687508 KP687469 KP687550 KP687589
Spain, Mallorca MA-Fungi 20615 KP687509 KP687470 KP687551 KP687590
Portugal, Estremadura MA-Fungi 31164 KP687510 KP687471 KP687552 KP687591
Spain, Córdoba MA-Fungi 55189 KP687528 KP687472 KP687553 KP687592
Spain, Cádiz MA-Fungi 59644 KP687511 KP687473 KP687554 KP687593
Spain, Madrid MA-Fungi 87325 KF988412 KF988540 KF988675 KF988808
Spain, Canary Islands Ribes 301207-43 KP687512 KP687474 KP687555 KP687594
Spain, Guadalajara Zamora 276 KP687513 KP687475 KP687556 KP687595
G. ovalisporum Bolivia, Concepción MA-Fungi 47184 KF988411 KF988539 KF988674 KF988805
Argentina, Salta MA-Fungi 86670 – KP687476 KP687557 –
Brazil, Rio Grande do Norte UFRN-Fungos 229 KP687514 – – –
G. papinuttii Argentina, Santiago del Estero MA-Fungi 83764 KF988380 KF988502 KF988637 KF988772
Argentina, Santiago del Estero MA-Fungi 86912 KP687515 KP687477 KP687558 KP687596
G. parvistriatum Spain, Madrid MA-Fungi 69583 JN943160 JN939560 JN991291 KF988806
Spain, Madrid Zamora 272 JN943162 JN939572 JN991283 KF988807
Spain, Madrid Zamora 285 JN943161 JN939571 JN991282 KP687597
G. pectinatum Spain, Lugo MA-Fungi 28156 KP687516 KP687478 KP687559 KP687598
Belgium MA-Fungi 43295 KP687517 KP687479 KP687560 –
Spain, Burgos MA-Fungi 75533 KP687518 KP687480 KP687561 –
Japan, Ibaraki TNS TKG-GE-70901 JN845111 JN845229 – JN845353
Japan, Ibaraki TNS TKG-GE-91201 JN845112 JN845230 – JN845354
Sweden, Uppland UPS F-149330 KP687519 KP687481 KP687562 KP687599
Sweden, Gotland UPS F-560803 KF988413 KF988541 KF988676 –
Spain, Huesca Zamora 290 KP687520 KP687482 KP687563 KP687600
Spain, Gerona Zamora 292 KP687521 KP687483 KP687564 KP687601
G. cf. plicatum Argentina, Buenos Aires MA-Fungi 83774 KF988415 KF988543 KF988678 KF988810
Argentina, Entre Ríos MA-Fungi 87322 KP687522 KP687484 KP687565 KP687602
Tanzania, Iringa UPS F-09935 KF988414 KF988542 KF988677 KF988809
Table 1   (cont.)
Taxon Country and state /province Herbarium voucher GenBank accession numbers
ITS LSU rpb1 atp6
G. quadrifidum Sweden, Uppland MA-Fungi 86671 KF988422 KF988550 KF988685 KF988817

USA, Colorado MICH 72512 KF988423 KF988551 KF988686 KF988818
Sweden, Södermanland S F-45993 JN845119 JN845237 – JN845361
Spain, Orense Zamora 139 KP687523 KP687485 KP687566 KP687603
Spain, Huesca Zamora 170 KF988421 KF988549 KF988684 KF988816
Spain, Cuenca Zamora 300 KP687524 KP687486 KP687567 KP687604
G. smithii Argentina, Córdoba MA-Fungi 83783 KF988442 KF988575 KF988710 KF988841
G. aff. smithii Australia, New South Wales CANB 748746 KP687525 KP687487 KP687568 KP687605
G. striatum Sweden, Uppland MA-Fungi 86672 KF988443 KF988577 KF988712 KF988843
Sweden, Närke S F-46074 JN845116 JN845233 – JN845358
Sweden, Uppland S F-74732 JN845117 JN845234 – JN845359
Spain, Valladolid Zamora 257 JN943164 JN939557 JN991288 KF988842
G. aff. striatum Mexico, Baja California AH 18521 – KP687530 KP687569 KP687608
G. tenuipes Australia, Australian Capital Territory CANB 738350 KP687526 KP687488 KP687570 KP687609
Australia, Australian Capital Territory CANB 775658 KP687527 KP687489 KP687571 KP687610
Australia, Victoria MEL 2096557 – DQ218602 – DQ218889
G. thanatophilum USA, Wisconsin MICH 72012 KF988364 KF988486 KF988621 KF988756
USA, Wisconsin MICH 72014 KF988365 KF988487 KF988622 KF988757
Geastrum sp. Japan, Aomori TNS TKG-GE-91002 JN845118 JN845236 – JN845360
Extraction Kit (QIAGEN, Germany). Sequencing was performed and length preset to 10 7 generations, sampling every 100th
by Macrogen (The Netherlands). tree. The analysis was automatically stopped when the aver-
DNA sequences were edited with Sequencher 4.1.4 (Gene age standard deviation across runs dropped below 0.005.
Codes, USA), primarily aligned using the FFT-NS-i strategy of Convergence was additionally assessed using Tracer v. 1.5
MAFFT (Katoh et al. 2002), and manually adjusted with BioEdit (Rambaut et al. 2013), by checking that the values of effective
v. 7.0 (Hall 1999), except the atp6 region that was directly sample size (ESS) for each parameter were ≥ 200, AWTY (Ny-
aligned with BioEdit reversely transcribed to protein format. lander et al. 2008) was used to discard lack of convergence by
Ambiguously aligned parts of ITS were removed with Gblocks visual inspection of the plots obtained from the sampled trees.
v. 0.91b (Castresana 2000), keeping default settings but allow- The first 25 % of the analysis was discarded as burn-in, and
ing all gap positions when not ambiguous. The remaining indels the 50 % majority-rule tree with branch lengths and posterior
were coded with FastGap v. 1.2 (Borchsenius 2007), using the probabilities (pp) was calculated from the remaining trees. In
simple indel coding method of Simmons & Ochoterena (2000), a preliminary analysis, similar or even more severe problems
in a separate binary data subset. Datasets are available in noted by Zamora et al. (2014) concerning convergence and
TreeBASE (TB2:S15972). overestimation of branch lengths (Brown et al. 2010, Marshall
2010) were detected in the present study. However, the se-
Maximum likelihood (ML) and Bayesian inference (BI) approach- lection of an appropriate exponential prior (1/λ) for obtaining
es were used for phylogenetic reconstructions. Conflicts among reasonable branch length estimates is often not obvious and
datasets were detected performing maximum parsimony boot- several trials may be required (Ekman & Blaalid 2011). To avoid
strap analyses of each region and identifying if a significantly this, new and less informative priors have been implemented
supported clade (bootstrap value ≥ 70 %, Hillis & Bull 1993) in MrBayes v. 3.2.2 (Rannala et al. 2012, Zhang et al. 2012).
from one DNA region is contradicted by another significantly We used a uniform compound Dirichlet prior ‘brlenspr = un-
supported clade from other regions. These preliminary boot- constrained : gammadir (1,1,1,1)’, obtaining rather reasonable
strap analyses were performed using the ‘fast’ stepwise-addition branch length estimates, and therefore this last analysis is the
bootstrap implemented in PAUP* v. 4.0b10 (Swofford 2003), one that will be shown and discussed.
with 1 000 non-parametric bootstrap replicates (Felsenstein
1985). Datasets were concatenated when no conflict was de- Relative strength of branch support values follows the scale
of Lutzoni et al. (2004). Phylogenetic trees were drawn using
tected.
FigTree v. 1.3 (Rambaut 2007).
ML analysis was done in GARLI v. 2.0 (Zwickl 2006), using the
following partitions: ITS1, 5.8S, ITS2, LSU, rpb1, atp6, and Morphological analyses
coded indels. The GTR+I+Γ model was used for each DNA
General methodology for collecting morphological data fol-
subset, and the Mk model for the binary subset (Lewis 2001).
lowed Sunhede (1989), Calonge (1998), and Zamora et al.
The analysis was repeated twice starting from random trees.
(2013). Macromorphological characters were measured in dried
For assessing branch supports, 1 000 non-parametric bootstrap
basidiomata. Micromorphological characters were measured
replicates were performed under the thorough bootstrap option
in 5 % KOH solution or in Hoyer’s medium. Basidiospore
of RAxML v. 7.4.2 (Stamatakis 2006), using the mentioned par
measurements included the ornamentation following Sunhede
titions and default settings of searching parameters.
(1989). Samples for scanning electron microscopy (SEM)
Bayesian inference was performed using the Metropolis Cou- were air dried, coated with pure gold in a Balzers SCD 004
pled Markov Chain Monte Carlo (MC 3) strategy implemented in sputter coater, and observed with a Hitachi S-3000N SEM. In
MrBayes v. 3.2.2 (Ronquist et al. 2012). DNA evolution models some cases, fragments of the endoperidium were previously
were selected with jModelTest v. 2.1 (Darriba et al. 2012), using washed with absolute ethanol to remove the excess of basidio
the Akaike Information Criterion (AIC). The F81 model was used spores. For old collections and some type material, already
for the binary subset of coded indels. Four parallel runs were detached fragments were used in order to avoid damaging
executed, each one starting with a random tree, with 6 chains, of the specimens. Calcium oxalate crystals were assigned to
COD or COM according to their habit (Frey-Wyssling 1981, the first two discriminant functions. The chi-square statistic
Horner et al. 1995). of Wilks’ lambda test (Stevens 1996) was used to assess
Continuous and discrete quantitative variables were measured the discriminatory capabilities of the discriminant functions,
on mature and dried herbarium basidiomata. Two morphometric i.e., if the functions differentiate the groups significantly.
approaches were used to test the utility of these characters in This set of multivariate analyses was performed in SPSS
establishing differences among the studied taxa. Statistics 22 (IBM Corp.). Finally, Cohen’s kappa and Z
test of significance were calculated according to Titus et al.
i) Most of the included species were more or less easily distin-
(1984), to evaluate whether there is agreement between
guished from the morphologically closest relatives according
the expected and the obtained classification of cases, and
to several qualitative and quantitative morphological traits.
if such agreement may be a product of chance, having into
For the newly proposed species, each quantitative character
account the sample sizes.
that may allow their distinction respect to the morphologically
b) The six most discriminant characters of each group were
most similar taxa was tested through an ANOVA, considering
represented as boxplots and analysed as in (i), i.e., first
p < 0.001 as significance value. When more than two taxa
through an ANOVA and, when significant differences found,
were involved, after detecting if there were significant dif-
using the Tukey’s HSD posthoc test to identify what means
ferences with the ANOVA, the Tukey’s honestly significance
were significantly different to others.
difference (Tukey’s HSD) posthoc test was used to detect
those means significantly different (p < 0.001) to each other. The following 13 morphological characters were measured: ba-
Measurements for each character were represented as box- sidiospore diameter (BASDIA), ornamentation height (ORN),
plots. These analyses and graphics were done using R (R maximum ornamentation height (ORNMAX), maximum dia
Development Core Team 2008). This approach was used for meter of the capillitial hyphae (CAP), maximum diameter of
comparing G. benitoi with G. arenarium, G. britannicum with isolated or twined, bipyramidal mesoperidial crystals of COD
G. quadrifidum and G. leptospermum, and G. kuharii and (CODCR), maximum diameter of mesoperidial crystalline ag-
G. thanatophilum with each other and with G. coronatum. gregates of COM (COMCR, only for the G. minimum group),
When enough material was available, up to 50 basidiomata diameter of the exoperidium not forced in horizontal posi-
of each species were used to measure macromorphologi- tion (apparent exoperidial diameter, EXAP), diameter of the
cal characters, the largest mesoperidial crystals, and the exoperidium when extended or forced in horizontal position
broadest capillitial hyphae. For basidiospore features, 100 (real exoperidial diameter, EXEXT), number of exoperidial rays
measurements were taken for G. arenarium, G. britannicum, (RAYS), diameter of the endoperidial body (END), stalk length
G. kuharii, G. leptospermum, and G. thanatophilum, and 200 (STL), stalk width (STW, largest diameter in the middle part),
measurements for G. benitoi, G. coronatum, and G. qua- and number of peristome folds (PER, only for the G. glauce
drifidum, because the amount of material of these last three scens and the G. pectinatum groups). In addition, the stalk
species was much higher. height /stalk width ratio (STL /W) was calculated as an index
of the robustness of the stalk. For basidiospore characteristics,
ii) For three species groups that are particularly difficult to
10 – 30 measurements were recorded per basidioma. Data
distinguish by morphology or have been consistently mis
used for DFAs are the mean values per basidioma rounded
interpreted in the literature, we performed multivariate
with a precision of 0.1 µm. For ANOVA analyses and boxplots
analyses on a wider dataset of quantitative morphological
representations, all measurements were considered to include
features. The three species groups so analysed were the
the whole variation observed.
‘G. minimum group’ (G. austrominimum, G. calceum s.l.,
G. granulosum, and G. marginatum), the ‘G. glaucescens
group’ (G. glaucescens, G. papinuttii, and G. parvistriatum), RESULTS
and the ‘G. pectinatum group’ (G. meridionale, G. pecti
natum, G. plicatum, and G. tenuipes). Thus, multivariate Molecular results
analyses were used to detect and evaluate putative useful A total of 160 DNA sequences were newly generated in this
morphological characters for distinguishing some of the study (Table 1). The concatenated matrix had 3 782 characters
clades (putative taxa) found in the previous phylogenetic (235 ITS1, 155 5.8S, 197 ITS2, 987 LSU, 1 184 rpb1, 707 atp6,
analyses. A priori assignation of specimens to particular and 317 coded indels), of which 2 365 were constant and 1 417
species was done by using a combination of all available variable. Maximum likelihood analyses recovered two trees with
data sources (morphology, DNA sequences, ecology, and lnL1 = -22178.0906 (best) and lnL2 = -22178.0931, both with a
chorology). Geastrum calceum s.l. samples were treated similar topology. The ML tree of the first replicate (the one with
as if they were a single species for morphological analyses the best likelihood score) is shown in Fig. 1.
due to the small sample size. The analysis with jModeltest yielded the following nucleotide
The suitability of the data for multivariate analyses was evalu- substitution models: HKY+Γ for ITS1, K80 for 5.8S, GTR+Γ for
ated for each analysed dataset under the Kaiser-Meyer-Olkin ITS2, LSU, and rpb1, and GTR+I+Γ for atp6. Bayesian MC3
measure of sampling adequacy (Kaiser 1974), calculated with runs were automatically halted after 3 315 000 generations.
the ‘rela’ package (Chajewski 2009) in R, and the Bartlett’s Best likelihood states for each run were lnL1 = -22292.50,
test of sphericity (Bartlett 1937), calculated with the ‘psych’ lnL2 = -22307.86, lnL3 = -22326.66, lnL4 = -22352.58. Potential
package (Revelle 2014), also in R. To avoid problems with Scale Reduction Factor values for model parameters were all
multicollinearity, the correlation matrix between each variable between 1.000 and 1.002. The topology of the 50 % majority
was calculated in SPSS Statistics 22 (IBM Corp.), and variables rule consensus tree is very similar to that of the ML tree, and
with a regression coefficient ≥ 0.95 were identified. One variable then only pp values are indicated on branches of the ML tree
of each pair with regression coefficient ≥ 0.95 was excluded in (Fig. 1).
the subsequent analyses. The ingroup (sect. Geastrum) of the ML tree (Fig. 1) is divided
Multivariate analyses were performed in two steps. into five strongly supported (bs ≥ 81 %, pp = 1.00) main clades,
a) The groups recognized in the phylogenetic analyses were considered as different subsections, which have been named
tested using a linear discriminant function analysis (DFA). from the base to the top of the tree as G. subsect. Arenaria,
Samples were graphically represented as score plots, using G. subsect. Hungarica, G. subsect. Quadrifida, G. subsect.
G. pectinatum TNS TKG-GE-91201

G. pectinatum TNS TKG-GE-70901
G. pectinatum MA-Fungi 43295
G. pectinatum Zamora 290
G. pectinatum Zamora 292
* G. pectinatum UPS F-149330 Epitype
G. pectinatum UPS F-560803
48/1 G. pectinatum MA-Fungi 28156
G. pectinatum MA-Fungi 75533
G. meridionale Zamora 276
G. meridionale MA-Fungi 87325 Holotype
G. meridionale Ribes 301207-43
G. meridionale MA-Fungi 55189
45/0.97 * G. meridionale AVM 2708
G. cf. biplicatum TNS TKG-GE-91003
G. cf. biplicatum TNS TKG-GE-91102
* G. cf. biplicatum TNS Sakamoto 191
* 64 G. cf. biplicatum TNS Sakamoto 182 subsect.
1 G. tenuipes CANB 738350 Sulcostomata
* G. tenuipes MEL 2096557
G. tenuipes CANB 775658
G. cf. plicatum MA-Fungi 87322
* G. cf. plicatum MA-Fungi 83774
G. cf. plicatum UPS F-09935
99 G. parvistriatum Zamora 272
1 * G. parvistriatum Zamora 285
64 G. parvistriatum MA-Fungi 69583 Holotype
1 * G. papinuttii MA-Fungi 86912
* G. papinuttii MA-Fungi 83764 Holotype
* G. glaucescens MA-Fungi 83762
G. glaucescens MA-Fungi 83763
G. striatum S F-74732
* G. striatum S F-46074
73/0.86
G. striatum Zamora 242
G. striatum MA-Fungi 86672
* G. striatum Zamora 257
98
G. striatum Zamora 251
1 G. aff. striatum AH 18521
G. coronatum Zamora 266
sect. Geastrum
* G. coronatum PRM 842868 (G. coronatum var. moelleri Holotype) subsect.
* G. coronatum MICH 28567
81 * G. aff. smithii CANB 748746 Geastrum
1 G. smithii MA-Fungi 83783
* G. thanatophilum MICH 72014
G. thanatophilum MICH 72012 Holotype
G. marginatum MA-Fungi 48129 Epitype
G. marginatum MA-Fungi 86669
G. marginatum ERRO 2012112609
* G. marginatum PRM 842884 (G. minimum var. fumosicollum Holotype)
G. marginatum MICH 28119b
89 G. quadrifidum MA-Fungi 86671
* G. quadrifidum Zamora 170
1
G. quadrifidum Zamora 300
51/0.98 G. quadrifidum Zamora 139
* G. quadrifidum S F-45993
G. quadrifidum MICH 72512
G. kuharii MA-Fungi 86914 subsect.
49/0.95 * G. kuharii MA-Fungi 83795 Holotype
G. kuharii MA-Fungi 86913 Quadrifida
84 G. cf. calceum2 UFRN-Fungos 723
G. calceum s.l.
G. cf. calceum1 MA-Fungi 83761
91 1
G. granulosum Zamora 191
1 G. granulosum K(M) 154623
G. granulosum MA-Fungi 69175
* G. granulosum Sunhede 7746
G. granulosum MICH 72010
95 G. granulosum MICH 28210
* 1 54/0.97 G. granulosum MICH 28119a
G. austrominimum MEL 2292062
96 * G. austrominimum CANB 748741 Holotype
1 G. austrominimum MEL 2358047
Geastrum sp. TNS TKG-GE-91002
G. hungaricum Sunhede 5993
G. hungaricum GB MJ9317 subsect.
* G. hungaricum GB MJ8915 Hungarica
G. hungaricum Zamora 611
G. benitoi Zamora 76
94
* G. benitoi MA-Fungi 68191
G. benitoi MA-Fungi 87324 Holotype
85 1 G. arenarium MA-Fungi 83760
1 G. ovalisporum UFRN-Fungos 229 subsect.
*
* G. ovalisporum MA-Fungi 47184 Holotype Arenaria
G. ovalisporum MA-Fungi 86670
G. britannicum K(M)79617 Holotype
* G. britannicum K(M)99914
G. britannicum K(M)60288
* G. fornicatum Zamora 255
G. fornicatum MA-Fungi 30749 outgroup (sect. Fornicata)
0.03
Fig. 1   Maximum likelihood phylogenetic tree of Geastrum sect. Geastrum. Numbers above branches indicate maximum likelihood bootstrap (bs) values,
and numbers below branches indicate Bayesian posterior probability (pp) values. Asterisks (*) represent branches with bs = 100 % and pp = 1. Only support
values above the species level are indicated. Type specimens are marked in bold.
Geastrum, and G. subsect. Sulcostomata. Geastrum subsect. bipyramidal crystals of COD (Fig. 2a, b), and crystalline ag-
Arenaria is composed by three strongly supported subclades gregates of COM (Fig. 2c). Bipyramidal crystals of COD can
(bs = 100 %, pp = 1.00), considered as three different spe- be isolated (Fig. 2a) to heavily twined (Fig. 2b), and were
cies: G. britannicum, G. ovalisporum, and G. benitoi, plus one found in all species, although they were normally scarce in
rather isolated specimen of G. arenarium. Geastrum subsect. G. hungaricum. In G. benitoi, these bipyramidal crystals were
Hungarica is composed by four specimens of a single spe- often present in the form of bipyramidal prisms, with conspicu-
cies, G. hungaricum. Geastrum subsect. Quadrifida is formed ous faces (Fig. 9b4), easily visible under the light microscope,
by six strongly supported subclades (bs ≥ 84 %, pp = 1.00), while in the other species mostly bipyramids or bipyramidal
distinguished as G. austrominimum, G. granulosum, G. cal prisms with hardly distinguished or less conspicuous faces
ceum s.l., G. kuharii, G. quadrifidum, and G. marginatum. The were present. Crystalline aggregates of COM were formed by
G. calceum s.l. subclade is formed by two specimens, G. cf. numerous thin scales (Fig. 2c); although often rounded, they
calceum1 (MA-Fungi 83761) and G. cf. calceum2 (UFRN-Fun- can be fused producing elongated forms. These crystalline ag-
gos 723), with notable differences in sequence data. Geastrum gregates have been only seen in species of Geastrum subsect.
subsect. Geastrum groups two strongly supported subclades Hungarica and Geastrum subsect. Quadrifida, although they are
(bs = 100 %, pp = 1.00), namely G. coronatum and G. thana- normally rare or indistinct in G. calceum s.l. and G. quadrifidum.
tophilum, plus two specimens, G. smithii (MA-Fungi 83783) Geastrum granulosum and G. marginatum showed a broad va-
and G. aff. smithii (CANB 748746), each one placed in well- riation of crystalline aggregates, that were normally scarce
separated branches, with notable differences in sequence and not well-developed, but sometimes abundant and big. The
data. Geastrum subsect. Sulcostomata includes nine strongly mesoperidium of G. coronatum sometimes lacked any kind of
supported subclades (bs = 100 %, pp = 1.00), recognized as crystalline matter.
the species G. striatum, G. glaucescens, G. papinuttii, G. par- Rhizomorph crystal morphology largely agreed with the types
vistriatum, G. cf. plicatum, G. tenuipes, G. cf. biplicatum, G. meri- recorded by Zamora et al. (2013). Bipyramidal crystals of COD,
dionale, and G. pectinatum, plus one specimen placed in a that were the most common type, were found in G. austromini
well-separated branch under the name G. aff. striatum. Finally, mum, G. benitoi, G. granulosum, G. kuharii, G. meridionale,
the specimen TNS TKG-GE-91002 is not included in any sub- G. ovalisporum, G. papinuttii, G. parvistriatum, G. quadrifidum,
section, and it is placed between G. subsect. Hungarica and G. striatum, and G. thanatophilum, forming rose-like aggregates
G. subsect. Quadrifida. or sometimes grouped on cystidioid-like cells (Fig. 2d). Irregular
horn-like crystals of COM may be found also in G. meridionale,
Morphological results G. ovalisporum, G. parvistriatum, and G. striatum. In G. coro
Scanning electron microscopy of the mesoperidial crystalline natum, thin horn-like crystals of COM, grouped in arachnoid
matter showed two well-differentiated morphological types, structures, were the dominant type (Fig. 2e). Novel in this study
Fig. 2   Mesoperidial and rhizomorph calcium oxalate crystals. a – c. Mesoperidial crystals: a. G. marginatum (MA-Fungi 86669) single bipyramidal crystal
of COD; b. G. kuharii (MA-Fungi 86913) twined bipyramidal crystals of COD; c. G. kuharii (MA-Fungi 86913) crystalline aggregate of COM scales. —
d – f. Rhizomorph crystals: d. G. parvistriatum (Zamora 539) cystidioid cell covered by bipyramidal crystals of COD, some grouped in rose-like aggregates;
e. G. coronatum (Zamora 484) arachnoid aggregate of thin horn-like COM crystals; f. G. leptospermum (lectotype) oblique prisms of COM grouped in stellate
aggregates. — Scale bars = 10 µm.
Geastrum arenarium / Geastrum benitoi
Pseudoparenchymaous cell walls

Basidiospore diameter
5.5
0.6
2.0
Ornamentation height
5.0
0.5
1.5
4.5
0.4
1.0
4.0
0.3
3.5
0.2
0.5
ARE BEN ARE BEN ARE BEN
a p<0.001* b p<0.001* c p<0.001*
Geastrum britannicum / Geastrum leptospermum / Geastrum quadrifidum

0.8
COD crystal diameter

6.0
70
0.6
5.0
50
0.4
4.0
30
3.0
0.2
10
BRI LEP QUA BRI LEP QUA BRI LEP QUA
d p<0.001*: BRI-LEP, BRI-QUA, LEP-QUA e p<0.001*: BRI-LEP, BRI-QUA, LEP-QUA f p<0.001*: BRI-QUA
p>0.001: BRI-LEP, LEP-QUA
0.5 1.0 1.5 2.0 2.5 3.0

14
Endoperidial diameter
Exoperidial diameter
50
12
Stalk height
40
10
30
8
20
6
4
10
BRI LEP QUA BRI LEP QUA BRI LEP QUA

g p<0.001*: BRI-LEP, BRI-QUA, LEP-QUA h p<0.001*: BRI-LEP, LEP-QUA i p<0.001*: BRI-LEP, BRI-QUA, LEP-QUA
p>0.001: BRI-QUA
Fig. 3   Boxplots representation of selected morphological characters for distinguishing new species in Geastrum subsect. Arenaria, and significance indices
of ANOVA or Tukey’s HSD tests. a – c: G. arenarium and G. benitoi. a. Basidiospore diameter; b. ornamentation height; c. thickness of the pseudoparenchyma-
tous cell walls. – d–i: G. britannicum, G. leptospermum, and G. quadrifidum. d. Basidiospore diameter; e. ornamentation height; f. maximum diameter of COD
mesoperidial crystals; g. exoperidial diameter; h. endoperidial diameter; i. stalk height. Species names abbreviated as: ARE = G. arenarium, BEN = G. benitoi,
BRI = G. britannicum, LEP = G. leptospermum, QUA = G. quadrifidum.
Geastrum coronatum / Geastrum kuharii / Geastrum thanatophilum

7.0

Capillitium diameter
20 40 60 80 100 120
12
6.5
10
6.0
8
5.5
5.0
6
4.5
COR KUH THA COR KUH THA COR KUH THA

a p<0.001*: COR-KUH, COR-THA, KUH-THA b p<0.001*: COR-KUH, COR-THA c p<0.001*: COR-KUH, COR-THA
p>0.001: KUH-THA p>0.001: KUH-THA
Endoperidial diameter
35
80 100 120
5
30
Stalk height
4
25
20
3
60
15
2
40
10
20
COR KUH THA COR KUH THA COR KUH THA

d p<0.001*: COR-THA e p<0.001*: COR-THA f p<0.001*: COR-THA
p>0.001: COR-KUH, KUH-THA p>0.001: COR-KUH, KUH-THA p>0.001: COR-KUH, KUH-THA
Fig. 4   Boxplots representation of selected morphological characters for distinguishing morphologically similar new species in Geastrum subsect. Geastrum
(G. thanatophilum) and Geastrum subsect. Quadrifida pro parte (G. kuharii ), and significance indices of Tukey’s HSD tests. a. Basidiospore diameter; b. maxi-
mum diameter of the capillitial hyphae; c. maximum diameter of COD mesoperidial crystals; d. exoperidial diameter; e. endoperidial diameter; f. stalk height.
Species names abbreviated as: COR = G. coronatum, KUH = G. kuharii, THA = G. thanatophilum.
is the presence of stellate aggregates of oblique prisms of COM showed regression coefficients ≥ 0.95 in correlation matrices,
in G. leptospermum rhizomorphs (Fig. 2f). and therefore the whole datasets were used for DFAs. The
The three selected continuous variables for distinguishing be- chi-square test of Wilk’s lambda was significant for all discri-
tween G. arenarium and G. benitoi showed significant differ- minant functions (Table 2). In the G. minimum group dataset
ences (p < 0.001) (Fig. 3a – c). The walls of the pseudoparen- the percentage of correctly classified samples was more than
chymatous layer cells were significantly thicker in G. arenarium, 90 % for the four taxa studied. Cohen’s kappa value was > 0.85
while basidiospores were significantly smaller and with a less for all species, but in G. calceum s.l. the Z test showed not
marked ornamentation. significant results (Table 2). In the corresponding score plot,
a low overlap is present among the different groups (Fig. 5a).
Geastrum britannicum, G. leptospermum, and G. quadrifidum In the G. glaucescens group dataset 100 % of G. papinuttii
also showed significant differences in all morphological char- samples were correctly classified, but less than 80 % of the
acters selected (ANOVA p < 0.001) (Fig. 3d – i). Geastrum G. glaucescens and G. parvistriatum samples were correctly
leptospermum showed the smallest and the least ornamented classified. Cohen’s kappa value was 1.00 for G. papinuttii, but
basidiospores, G. quadrifidum had the biggest and the most < 0.65 for G. glaucescens and G. parvistriatum. The Z test was
ornamented ones, and in G. britannicum basidiospores were significant for all this values (Table 2). The score plot of this
intermediate between them. Tukey’s HSD test also showed group showed a wide overlap between G. glaucescens and
significant differences in all possible comparisons among the G. parvistriatum (Fig. 5b). In the G. pectinatum group dataset
three species (Fig. 3d, e). Geastrum britannicum showed the the percentage of correctly classified samples was 95 % for
biggest mesoperidial crystals of COD, while G. quadrifidum had G. meridionale and 100 % for G. pectinatum, G. plicatum, and
the smallest; G. leptospermum mesoperidial crystals showed G. tenuipes. Cohen’s kappa value was > 0.90 for all species and
an intermediate size. Tukey’s HSD test showed significant the Z test was significant (Table 2). The score plot, representing
differences only between G. britannicum and G. quadrifidum the first two discriminant functions, showed that G. plicatum
(Fig. 3f). Geastrum britannicum was also the species with the and G. tenuipes were clearly separated, while G. meridionale
largest basidiomata (exoperidial and endoperidial diameters, and G. pectinatum greatly overlap (Fig. 5c).
and stalk height), G. quadrifidum was somewhat smaller, and
The following characters were selected as the most discriminant
G. leptospermum was much smaller than both of them. Signifi
in the G. minimum group: basidiospore diameter, ornamentation
cant differences were found in Tukey’s HSD test for all possible
height, maximum ornamentation height, maximum diameter
comparisons of these characters except for the endoperidial
of mesoperidial COD crystals, maximum diameter of meso
diameter of G. britannicum and G. quadrifidum (Fig. 3g–i).
peridial COM aggregates, and exoperidial diameter (Table 2).
All the characters used for separating G. coronatum, G. kuharii, All of them showed significant differences among the included
and G. thanatophilum showed significant differences (ANOVA species (Fig. 6). Geastrum austrominimum had the biggest
p < 0.001) (Fig. 4). Geastrum coronatum had the largest basidio- mesoperidial aggregates of COM (Fig. 6e). Geastrum calceum
spores, G. kuharii the smallest, and in G. thanatophilum they showed the basidiomata with the largest size, represented as
were intermediate. Significant differences were found in Tukey’s the exoperidial diameter (Fig. 6f), and basidiospores with the
HSD test for all possible comparisons (Fig. 4a). The capillitium highest ornamentation, followed by G. marginatum (Fig. 6b, c).
of G. coronatum was significantly wider than in G. kuharii and The mesoperidial COD crystals of G. granulosum and G. cal
G. thanatophilum, while significant differences were not found ceum s.l. were the biggest, and those of G. marginatum were
for these two last taxa (Fig. 4b). Geastrum kuharii and G. tha the smallest (Fig. 6d). Significant differences were found in the
natophilum mesoperidial crystals of COD were rather similar basidiospore diameter, but measurements of this character also
between them and significantly bigger than those of G. corona showed a broad overlapping (Fig. 6a).
tum (Fig. 4c). Basidiomata of G. coronatum were the biggest, In the G. glaucescens group, DFA identified the following cha-
represented by both the exoperidial and endoperidial diameters, racters as the most discriminant: basidiospore diameter, orna-
and also showed the longest stalks, G. thanatophilum was the mentation height, maximum ornamentation height, capillitium
smallest and had the shortest stalks, and G. kuharii showed diameter, exoperidial diameter, and number of peristome folds
intermediate macromorphological characters between them; for (Table 2). Geastrum papinuttii had the smallest and less orna
these characters only comparisons between G. coronatum and mented basidiospores (Fig. 7a–c), as well as the thinnest ca-
G. thanatophilum showed significant differences (Fig. 4d–f). pillitial hyphae (Fig. 7d) and the smallest basidiomata, repre
The Kaiser-Meyer-Olkin measure of sampling adequacy was sented by the exoperidial diameter (Fig. 7e). The number of
> 0.7, and the Bartlett’s test of sphericity was significant for all peristome folds did not show significant differences in any
matrices used in multivariate analyses (Table 2). No variables comparison pairs (Fig. 7f). Significant differences between
Table 2   Results from DFAs indicating the number of basidiomata (N), Kaiser-Meyer-Olkin measure of sampling adequacy (KMO), Barlett’s test, Wilks’ lambda
test, significance of chi-square statistic of Wilks’ lambda, percentage of correctly classified samples, Cohen’s kappa and significance of Z test (* means
p < 0.001, ns means p > 0.001), and the six most discriminant variables selected for each group. AUS = G. austrominimum, CAL = G. calceum s.l., GRA =
G. granulosum, MAR = G. marginatum, GLA = G. glaucescens, PAP = G. papinuttii, PAR = G. parvistriatum, MER = G. meridionale, PEC = G. pectinatum,
PLI = G. plicatum, TEN = G. tenuipes.
Dataset N KMO Bartlett’s Wilks’ λ Sig. χ2 Wilks’ λ % correctly class. Cohen’s κ(sig.) Selected variables
G. minimum group AUS: 27 0.707 p < 0.001 DF1 to 3: 0.053 DF1 to 3: p < 0.001 AUS: 92.6 AUS: 0.883* BASDIA ORN
CAL: 5 DF2 to 3: 0.235 DF2 to 3: p < 0.001 CAL: 100 CAL: 1.000ns ORMAX CODCR
GRA: 89 DF3: 0.625 DF3: p < 0.001 GRA: 94.4 GRA: 0.911* COMCR EXEXT
MAR: 70 MAR: 95.7 MAR: 0.932*
G. glaucescens group GLA: 28 0.806 p < 0.001 DF1 to 2: 0.082 DF1 to 2: p < 0.001 GLA: 78.6 GLA: 0.633* BASDIA ORN
PAP: 16 DF2: 0.632 DF2: p < 0.001 PAP: 100 PAP: 1.000* ORNMAX CAP
PAR: 63 PAR: 79.4 PAR: 0.647* EXEXT PER
G. pectinatum group MER: 40 0.794 p < 0.001 DF1 to 3: 0.007 DF1 to 3: p < 0.001 MER: 95 MER: 0.923* BASDIA CAP
PEC: 21 DF2 to 2: 0.072 DF2 to 3: p < 0.001 PEC: 100 PEC: 1.000* CODCR STW
PLI: 11 DF3: 0.295 DF3: p < 0.001 PLI: 100 PLI: 1.000* STL/W PER
TEN: 13 TEN: 100 TEN: 1.000*
G. minimum group G. glaucescens and G. parvistriatum were only found in basidio

spore diameter, although boxplots showed a complete overlap
of measurements (Fig. 7a).
5
For the G. pectinatum group, the most discriminant morphologi-
cal characters were the basidiospore size, capillitium diameter,
maximum diameter of mesoperidial COD crystals, stalk width,
stalk length /stalk width ratio, and number of peristome folds
3
(Table 2). Basidiospores of G. plicatum were significantly
smaller than in the remaining species (Fig. 8a). The capillitium of
G. pectinatum was significantly wider than in the other species
(Fig. 8b). Geastrum tenuipes showed mesoperidial crystals of
DF 2
1 COD significantly larger than other species (Fig. 8c). Geastrum

meridionale had the thickest and stoutest stalks, followed by
G. pectinatum, while stalks of G. plicatum and G. tenuipes were
significantly thinner and more slender (Fig. 8d, e). Finally, the
-1 number of peristome folds was significantly higher in G. meri
dionale than in the other species (Fig. 8f).
TAXONOMY
-3
Geastrum section Geastrum

a -4,5 -2,5 -0,5
DF 1
1,5 3,5 5,5
Type. Geastrum coronatum Pers. (proposed as conserved type for
G. glaucescens group Geastrum by Zamora (2014)).
4
Key to subsections in Geastrum section Geastrum
1. Peristome always sulcate; mostly conical to narrowly coni-

cal; mesoperidium as a very thick, often farinose, layer of
2 pruina made up by generative hyphae and rather small
crystals (mostly < 20 µm diam, except G. tenuipes up to
55 µm diam) . . . . . . . . . . . . . . . G. subsect. Sulcostomata
DF 2
1. Peristome often fibrillose, rarely sulcate; mostly flat to broadly

conical; mesoperidium different . . . . . . . . . . . . . . . . . . . . 2
0 2. Basidiospores mostly 3.0–5.0 µm diam and inconspicuously
ornamented (ornamentation mostly 0.1– 0.5 µm high) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. subsect. Arenaria
2. Basidiospores mostly 4.5–7.0 µm diam and conspicuously
ornamented (ornamentation mostly 0.4 –1.0 µm high) . . . 3
-2
3. Exoperidium ± saccate, strongly hygrometric; pseudoparen-
chymatous layer very persistent, made up by thick-walled
(> 1.5 µm thick) hyphal cells; endoperidial body sessile to
nearly so . . . . . . . . . . . . . . . . . . . . . G. subsect. Hungarica
b -7,5 -5,0 -2,5 0,0 2,5 3. Exoperidium not saccate, not hygrometric; pseudoparen-
DF 1
chymatous layer evanescent, made up by thin-walled (mostly
G. pectinatum group ≤ 1.0 µm thick) hyphal cells; endoperidial body distinctly
stalked (stalk sometimes short) . . . . . . . . . . . . . . . . . . . . 4
4. Basidiospores with processes not covered with warts or
3 only with some irregular and inconstant warts; basidiomata
often delicate and slender, rarely stout; mesoperidium with
crystalline matter predominant over the generative hyphae
(sometimes crystals and hyphae both abundant), often with
1 crystalline aggregates of COM; peristome always fibrillose
. . . . . . . . . . . . . . . . . . . . . . . . . . . . G. subsect. Quadrifida
4. Basidiospores with processes densely covered by regular
and small rounded warts; basidiomata rather stout; mesoperi-
DF 2
-1
dium with generative hyphae predominant over the crystals
(sometimes only generative hyphae present), without crystal-
line aggregates of COM; peristome fibrillose to sulcate . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . G. subsect. Geastrum
-3
←
Fig. 5   Score plots of the G. minimum, G. glaucescens, and G. pectinatum
groups, using the two first discriminant functions from the discriminant func-
tion analyses. — a. G. minimum group: G. calceum s.l.; G. granulosum;
-5
G. marginatum; G. austrominimum. – b. G. glaucescens group: G.
c -4,0 -1,5 1,0 3,5 6,0 8,5 papinuttii; G. glaucescens; G. parvistriatum. – c. G. pectinatum group:
DF 1 G. tenuipes; G. plicatum; G. pectinatum; G. meridionale.
Geastrum minimum group (G. austrominimum / G. calceum / G. granulosum / G. marginatum)
1.2
Maximum ornamentation height

7.5
1.1
1.0
6.5
0.9
0.8
0.6
5.5
0.7
0.4
4.5
0.5
AUS CAL GRA MAR AUS CAL GRA MAR AUS CAL GRA MAR
a p<0.001*: all but AUS-CAL b p<0.001*: all but AUS-MAR c p<0.001*: AUS-CAL, CAL-GRA, GRA-MAR
p>0.001: AUS-CAL p>0.001: AUS-MAR p>0.001: AUS-GRA, AUS-MAR, CAL-MAR
COM aggregate diameter

200
60
80 100
50
150
40
100
60
30
40
50
20
20
10
0
AUS CAL GRA MAR AUS CAL GRA MAR AUS CAL GRA MAR
d p<0.001*: AUS-GRA, AUS-MAR, CAL-MAR, GRA-MAR e p<0.001*: AUS-CAL, AUS-GRA, AUS-MAR, GRA-MAR f p<0.001*: AUS-CAL, CAL-GRA, CAL-MAR
p>0.001: AUS-CAL, CAL-GRA p>0.001: CAL-GRA, CAL-MAR p>0.001: AUS-MAR, AUS-GRA, GRA-MAR
Fig. 6   Boxplots representation of the most discriminant characters retrieved by DFA for the G. minimum group, and significance indices of Tukey’s HSD
tests. a. Basidiospore diameter; b. ornamentation height; c. maximum ornamentation height; d. maximum diameter of COD mesoperidial crystals; e. maximum
diameter of COM mesoperidial crystal aggregates; f. exoperidial diameter. Species names abbreviated as: AUS = G. austrominimum, CAL = G. calceum s.l.,
GRA = G. granulosum, MAR = G. marginatum.
Geastrum glaucescens group (G. glaucescens / G. papinuttii / G. parvistriatum)

Maximum ornamentation height
1.2
7.0
1.1
1.0
6.0
0.9
0.8
0.6
5.0
0.7
0.4
4.0
0.5
GLA PAP PAR GLA PAP PAR GLA PAP PAR

a p<0.001*: GLA-PAP, GLA-PAR, PAP-PAR b p<0.001*: GLA-PAP, PAP-PAR c p<0.001*: GLA-PAP, PAP-PAR
p>0.001: GLA-PAR p>0.001: GLA-PAR
25
9
45
Peristome folds
8
20
35
7
15
25
6
10
5
15
GLA PAP PAR GLA PAP PAR GLA PAP PAR

d p<0.001*: GLA-PAP, PAP-PAR e p<0.001*: GLA-PAP, PAP-PAR f p>0.001: GLA-PAP, GLA-PAR, PAP-PAR
p>0.001: GLA-PAR p>0.001: GLA-PAR
Fig. 7   Boxplots representation of the most discriminant characters retrieved by DFA for the G. glaucescens group, and significance indices of Tukey’s HSD
tests. a. Basidiospore diameter; b. ornamentation height; c. maximum ornamentation height; d. maximum diameter of the capillitial hyphae; e. exoperidial
diameter; f. number of peristome folds. Species names abbreviated as: GLA = G. glaucescens, PAP = G. papinuttii, PAR = G. parvistriatum.
Geastrum subsect. Arenaria J.C. Zamora in Zamora et al., 3. Pseudoparenchymatous layer persistent, made up by more
Taxon 63, 3: 490. 2014 or less thick-walled (mostly 1.0–2.0 µm thick) hyphal cells;
exoperidium hygrometric; American species G. arenarium
Key to species in Geastrum subsection Arenaria 3. Pseudoparenchymatous layer rather evanescent, made up
by thin-walled (mostly ≤ 1.0 µm thick) hyphal cells; exoperi
1. Stalk stout, ≤ 1.0 mm high . . . . . . . . . . . . . . . . . . . . . . . . 2 dium not to falsely hygrometric; South European species
1. Stalk more or less slender, ≥ 1.3 mm high . . . . . . . . . . . . 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. benitoi
2. Exoperidium fornicate; largest mesoperidial crystals 40–60 4. Basidiospores mostly globose to subglobose, (3.5–)4.0–
µm diam; species growing among mosses on bark of living 4.5(–5.0) µm diam; ornamentation 0.3–0.5(–0.6) µm high;
trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. leptospermum exoperidium fornicate; Paleartic species . . G. britannicum
2. Exoperidium saccate to arched; largest mesoperidial crystals 4. Basidiospores mostly ovoid, 3.0–4.0 × 2.5–3.5 µm; orna-
10 – 35 µm diam; species growing in rather dry habitats and mentation 0.1–0.3 µm high; exoperidium arched; Neotropical
sandy soils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 species . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. ovalisporum
Geastrum pectinatum group (G. meridionale / G. pectinatum / G. plicatum / G. tenuipes)


14
50
8.0
40
12
7.0
30
10
6.0
20
8
10
6
5.0
MER PEC PLI TEN MER PEC PLI TEN MER PEC PLI TEN
a p<0.001*: MER-PLI, PEC-PLI, PLI-TEN b p<0.001*: MER-PEC, PEC-PLI, PEC-TEN c p<0.001*: MER-TEN, PEC-TEN, PLI-TEN
p>0.001: MER-PEC, MER-TEN, PEC-TEN p>0.001: MER-PLI, MER-TEN, PLI-TEN p>0.001: MER-PEC, MER-PLI, PEC-PLI
Stalk length/stalk width
60
8
Peristome folds
8
50
Stalk width
6
6
40
4
4
30
2
2
20
MER PEC PLI TEN MER PEC PLI TEN MER PEC PLI TEN
d p<0.001*: MER-PLI, MER-TEN, PEC-PLI, PEC-TEN e p<0.001*: MER-PLI, MER-TEN, PEC-PLI, PEC-TEN f p<0.001*: MER-PEC, MER-PLI, MER-TEN
p>0.001: MER-PEC, PLI-TEN p>0.001: MER-PEC, PLI-TEN p>0.001: PEC-PLI, PEC-TEN, PLI-TEN
Fig. 8   Boxplots representation of the most discriminant characters retrieved by DFA for the G. pectinatum group, and significance indices of Tukey’s HSD tests.
a. Basidiospore diameter; b. maximum diameter of the capillitial hyphae; c. maximum diameter of COD mesoperidial crystals; d. stalk width; e. stalk length /stalk
width ratio; f. number of peristome folds. Species names abbreviated as: MER = G. meridionale, PEC = G. pectinatum, PLI = G. plicatum, TEN = G. tenuipes.
Geastrum arenarium Lloyd, Bull. Lloyd Libr. Bot. 5: 28. 1902, part of this collection or explicitly excluding the non-conform
‘Geaster arenarius’ — Fig. 3, 9a species (Art. 9.17).
Type. Lectotype (designated here as a ‘second-step’ lectotype, MBT
198515): USA, Florida, Jupiter, Feb. 1895, H.C. Culbertson, Smithsonian Geastrum benitoi J.C. Zamora, sp. nov. — MycoBank
Institution 57292, BPI 704841!, excluding one basidioma of G. quadrifidum MB810499; Fig. 3, 9b
and one basidioma of Astraeus sp. (as A. hygrometricus) that are marked
separately. Type. Spain, Madrid, Villaviciosa de Odón, urbanización Campodón, under
Olea europaea, on sandy, siliceous soil, 12 Nov. 2011, B. Zamora & J.C.
Synoptic description (based on Sunhede (1986) and studied Zamora, Zamora 499, holotype MA-Fungi 87324!, isotypes in AH 45201! and
specimens) — Exoperidium 15 – 40 mm diam in horizontal po- UPS!
sition, saccate to arched when hydrated, with (6 –)7–10(–13) Etymology. The specific epithet is dedicated to Benito Zamora, father of
hygrometric rays. Endoperidial body 5 –19 mm diam, whitish to the first author, who has been helping him to collect Geastrum specimens
grey or brownish grey. Mesoperidium as a thin layer of pruina, during years of field trips.
sometimes inconspicuous. Largest mesoperidial crystals of
COD on the endoperidial surface 15–30 µm diam, bipyramidal. Diagnosis — Exoperidium (13–)15–35(–38) mm diam in hori-
zontal position, arched, with (6–)7–11(–13) often falsely hygro-
Peristome fibrillose, distinctly delimited. Stalk mostly 0.2–1.0
metric rays. Endoperidial body (4–)5–11(–13) mm diam, mostly
mm high, rarely indistinct, whitish. Basidiospores mostly glo-
greyish cream to brownish grey, rarely whitish. Mesoperidium
bose to subglobose, some ovoid, 3.5 – 4.5(– 5.0) µm diam, with
as a thin layer of pruina. Largest mesoperidial crystals of COD
(0.2–)0.3–0.4(–0.7) µm high warts, ornamentation verrucose.
on the endoperidial surface 18–35 µm diam, often bipyramidal
Pseudoparenchymatous layer persistent, of more or less thick-
prisms with well-developed faces, but also bipyramids. Peri
walled (mostly 1– 2 µm thick) cells.
stome fibrillose, well-delimited. Stalk 0.2–1.0 mm high, whit-
Ecology & Distribution — Originally described from sandy
ish to cream. Basidiospores mostly globose to subglobose,
soils of the ‘Temperate conifer forests’ biome of the Neartic
(3.5 –)4.0 – 5.0(– 5.5) µm diam, with 0.3 – 0.5(– 0.6) µm high
ecozone (Lloyd 1902). It has been also found in ‘Deserts and
warts, ornamentation verrucose. Pseudoparenchymatous layer
xeric shrublands’ and ‘Tropical and subtropical grasslands, of thin-walled (mostly ≤ 1 µm thick) cells. Rhizomorphs with
savannas and shrublands’ biomes of the Neartic (Bates 2004) bipyramidal crystals of COD, isolated or grouped in rose-like
and Neotropical (Kuhar et al. 2013) ecozones. Unconfirmed aggregates.
records have been also made from the Afrotropical (Bottomley
1948) and Australasian (Cunningham 1944) ecozones. Macroscopic characteristics — Unexpanded basidiomata
4–7 mm diam, subglobose, with a rounded apex or a flat umbo.
Additional specimens examined. Argentina, Catamarca, camino a ‘Cha
Exoperidium splitting in (6 –)7–11(–13) more or less equal
ñaritos’, entre mantillo, bajo jarilla y quebracho blanco, Chaco serrano árido,
2 Mar. 1994, M.M. Dios, G. Castro & L. Rodríguez, BAFC 33321; Córdoba, to unequal rays, (9–)10–30(–33) mm diam apparently, (13–)15–
Pocho, reserva Chancaní, cerca de la vivienda, bajo Prosopis chilensis, 35(–38) mm diam when forced in horizontal position, arched,
7 June 1993, L. Domínguez & E. Crespo 1333, CORD 1333; La Rioja, often falsely hygrometric. Mycelial layer thin, whitish to pale
Miranda, Prosopis sp., 27 Mar. 2008, L. Papinutti & G. Rolón, MA-Fungi cream, strongly intermixed with debris from the substrate, more
83760. – USA, Arizona, 7 mi. from Nogales, 11 Sept. 1941, Long & Stouffer or less strongly adhered to the fibrous layer, but sometimes
9634, LPS 29761; Arizona, 10 mi. from Nogales, along Tucson-Nogales peeling-off in some parts. Fibrous layer papyraceous to slightly
highway, 4 June 1930, Long 8304, LPS 29751; Arizona, Tucson, near Sabino coriaceous when denuded, whitish to pale-cream coloured.
Canyon, 4 June 1938, W.H. Long & Stouffer 9263, LPS 29763; Florida, locis
Pseudoparenchymatous layer pale cream to greyish cream,
arenosis, C.G. Lloyd 213, S.
not or only superficially cracked, < 0.5 mm thick when dried,
Nomenclatural notes — Sunhede (1986) firstly lectotypified about 1 mm thick when fresh, more or less evanescent. Endo
the species with BPI 704841, but without explicitly selecting a peridial body globose to subglobose, rarely irregular, (4–)5–
Fig. 9   Morphological characters in Geastrum subsect. Arenaria and Geastrum subsect. Hungarica. a. G. arenarium (MA-Fungi 83760 and S (C.G. Lloyd
213)); b. G. benitoi (holotype); c. G. britannicum (holotype); d. G. leptospermum (lectotype); e. G. ovalisporum (holotype); f. G. hungaricum (Zamora 611).
a1, b1, c1, d1, e1, f1, f2, basidiomata habit, f1 = f2 but wetted to show hygrometry of exoperidial rays; a2, b2, c2, d2, e2, detail of the peristome; a3, b3, c3,
d3, e3, detail of the stalk and apophysis; a4, b4, c4, d4, e4, f3, mesoperidial crystalline matter on the endoperidial surface, a4, b4, c4, d4, e4 show bipyramidal
crystals of COD, f3 shows crystalline aggregates of COM scales; a5, b5, c5, d5, e5, f4, basidiospores. — Scale bars: a1, b1, c1, d1, e1, f1, f2 = 5 mm; a2, b2,
c2, d2, e2, a3, b3, c3, d3, e3 = 2 mm; a4, b4, c4, d4, e4, f3 = 10 µm; a5, b5, c5, d5, e5, f4 = 2 µm.
11(–13) mm diam, greyish cream to whitish; endoperidial sur- Zamora, herb. Zamora 542; ibid., 15 Mar. 2014, B. Zamora & J.C. Zamora,
face glabrous or almost so, in newly expanded basidiomata herb. Zamora 615.
covered with very small, bipyramidal crystals, often in the form
of bipyramidal prisms with conspicuous faces, bipyramids also Geastrum britannicum J.C. Zamora, sp. nov. — MycoBank
present, gradually disappearing. Peristome fibrillose, mostly MB810500; Fig. 3, 9c
darker than the endoperidial surface, broadly conical to almost
Type. UK (England), Norfolk, Cockley Cley, in litter, Pinus sylvestris, 25
flat, 0.5 –1.5 mm high, well-delimited. Stalk present but often
Sept. 2000, J. Revett, holotype K(M) 79617!
very short, 0.2 –1.0 mm high, whitish. Apophysis absent or
poorly developed, concolorous with the endoperidium. Colu Etymology. The specific epithet refers to the only country where the spe-
mella intruding about 1/3 –1/2 into the glebal mass. Mature cies is known from.
gleba dark greyish brown. Diagnosis — Exoperidium 44–52 mm diam in horizontal posi-
Microscopic characteristics — Basidia narrowly ellipsoid to tion, fornicate, with 4 –7 non-hygrometric rays. Endoperidial
subcylindric or more or less lageniform, 15 – 23 × 4.5 – 6.5 µm, body 11–12 mm diam, pale to dark brown. Mesoperidium well-
with (3 –)4 –7(– 8) short sterigmata. Basidiospores mostly glo- formed, with abundant crystalline matter. Largest mesoperidial
bose to subglobose, a few ovoid, (3.5–)4.0–5.0(–5.5) µm diam, crystals of COD on the endoperidial surface 45–80 µm diam,
brownish to yellowish brown, with 0.3–0.5(–0.6) µm high brown, bipyramidal, often grouped in 60–120 µm diam rounded ag-
irregular warts, ornamentation verrucose. Broadest capillitial gregates. Peristome fibrillose, well-delimited. Stalk 2.0 – 3.0
hyphae 4.5 – 6.5 µm wide, aseptate, very rarely branched, nor- mm high, brownish. Basidiospores globose to slightly ovoid,
mally straight, thick-walled (2.0 – 3.0 µm thick), with narrow (3.5–)4–4.5(–5) µm diam, with (0.3–)0.4–0.5(–0.6) µm high
lumen, mostly visible; tips acute to rounded; surface covered warts, ornamentation verrucose. Pseudoparenchymatous layer
with debris or not. Endoperidium composed of 2.0 – 5.0(–6.0) of thin-walled (mostly ≤ 1 µm thick) cells.
µm wide, yellowish to yellowish brown, aseptate, mostly un-
branched, slightly sinuous, strongly intertwined, thick-walled Macroscopic characteristics — Unexpanded basidiomata
hyphae, lumen visible; protruding hyphae absent or very not seen. Exoperidium split in 4 –7 more or less equal to
sparse and almost indistinct. Peristomal hyphae 2.5 –7.5 µm unequal rays, 30–50 mm diam apparently, 44–52 mm diam
wide, light brown, aseptate, mostly unbranched, thick-walled when forced in horizontal position, fornicate when mature,
(1.0–2.5 µm thick), lumen visible, straight to somewhat sinu- not hygrometric. Mycelial layer comparatively thick, whitish to
ous, narrowing at base and apex, tips mostly acute to more or pale cream, strongly intermixed with debris from the substrate,
less rounded. Mesoperidium present on the endoperidium and separating from the fibrous layer and remaining as an irregular
pseudoparenchymatous layer surfaces, consisting of 18–35 mycelial cup attached to the tips of the rays. Fibrous layer more
µm diam, bipyramidal crystals of COD, intermixed with some or less coriaceous when denuded, whitish to cream coloured.
1.0 – 3.0 µm wide, hyaline, branched, thin-walled, clamped Pseudoparenchymatous layer pale cream to ochraceous cream,
hyphae. Pseudoparenchymatous layer of thin-walled (mostly frequently cracked, less than 1 mm thick in dry state, peeling
≤ 1 µm thick), hyaline to yellowish cells, variable in shape off from the fibrous layer and not very persistent (absent in old
and size, about 12 –75 × 10 – 32 µm. Fibrous layer with 1.5–5 basidiomata). Endoperidial body subglobose to ovoid, some-
µm wide, hyaline to very pale yellowish, aseptate, straight or times irregular, 11–12 mm diam, pale to dark brown; endoperidial
slightly sinuous, intertwined, mostly unbranched, comparatively surface glabrous or almost so, in newly expanded basidiomata
thick-walled (0.5 – 2.0 µm thick) hyphae, lumen more or less densely covered with big, bipyramidal crystals, often forming
visible. Mycelial layer double-layered; inner layer consisting of rounded aggregates, remaining long time and gradually disap-
1.5–3 µm wide, strongly glued together, more or less hyaline, pearing. Peristome fibrillose but with up to 10 inconspicuous and
branched, thin-walled and clamped hyphae; outer layer with poorly developed folds near the centre, mostly darker than the
1–3.5 µm wide, hyaline to somewhat yellowish, aseptate, rarely endoperidial surface, broadly conical to almost flat, 1.0–1.5 mm
branched, comparatively more or less thick-walled (0.5–1.5 µm high, sharply delimited. Stalk present, well-developed, 2.0–3.0
thick) hyphae, lumen hardly visible. Rhizomorphs covered with mm high, brownish. Apophysis well-defined, concolorous or
rose-like aggregates of bipyramidal crystals. slightly lighter than the endoperidium. Columella intruding about
Ecology & Distribution — It is only known from sandy, sili 1/2 into the glebal mass. Mature gleba dark greyish brown.
ceous soils of the Iberian Peninsula, which is part of the ‘Medi- Microscopic characteristics — Basidia not seen. Basidio
terranean forests, woodlands and scrub’ biome of the Paleartic spores globose to slightly ovoid, (3.5–)4.0–4.5(–5.0) µm diam,
ecozone. brownish to yellowish brown, with (0.3–)0.4–0.5(–0.6) µm high
brown warts, ornamentation verrucose, hilar appendix 0.5–0.8
Additional specimens examined (paratypes). Spain, Badajoz, Campa
µm long. Broadest capillitial hyphae 3.5–9.5 µm wide, aseptate,
nario, Badija, open area, sandy, siliceous soil, 2 Nov. 2013, M. Gordillo, herb.
Zamora 615; Jaén, Santuario de la Virgen de la Cabeza, en jaral, 30 Dec.
very rarely branched, normally straight, thick-walled (1.5–4.5
1984, A.G. Buendía, MA-Fungi 8106; Madrid, Casa de Campo, 6 Dec. 1985, µm thick), with narrow lumen, mostly visible; tips acute to
M. Jeppson, MA-Fungi 16940; Madrid, Colmenarejo, río Aulencia, 30TVK1582, rounded; surface often covered with debris. Endoperidium com-
635 m, en pradera, 20 Oct. 2001, F. Prieto GP292, MA-Fungi 68191; ibid., posed of 1.5–5.0 µm wide, pale yellowish to yellowish brown,
suelo arenoso, casi desnudo, en las proximidades de Retama sphaero aseptate, mostly unbranched, slightly sinuous, strongly inter-
carpa y Quercus rotundifolia, 28 Jan. 2002, F. Prieto, herb. Zamora 208; twined, thick-walled hyphae, lumen visible; protruding hyphae
ibid., 20 Feb. 2002, F. Prieto & Á. González, herb. Zamora 209; ibid., 4 Jan.
not seen. Peristomal hyphae 3.0–5.5 µm wide, brown to dark
2007, F. Prieto & J.C. Zamora, herb. Zamora 210; Madrid, Villaviciosa de
Odón, urbanización Campodón, bajo olivos en un antiguo olivar abandonado,
brown, aseptate, mostly unbranched, thick-walled (1.0–2.5 µm
17 Mar. 2001, J.C. Zamora, MA-Fungi 53523 (duplo herb. Zamora 13); ibid., thick), lumen mostly visible, often sinuous, narrowing at base
under Olea europaea, on sandy, siliceous soil, 27 Feb. 2005, J.C. Zamora, and apex, tips mostly rounded, a few acute; abundant debris
herb. Zamora 76; ibid., 14 Dec. 2005, B. Zamora & J.C. Zamora, herb. present among hyphae. Mesoperidium present on the endo
Zamora 124, ibid., 20 Oct. 2006, J.C. Zamora, herb. Zamora 180; ibid., 3 Feb. peridium and pseudoparenchymatous layer surfaces, consisting
2007, B. Zamora & J.C. Zamora, herb. Zamora 214; ibid., 8 Dec. 2008, of rather large, bipyramidal crystals, 20 – 80 µm diam (the
B. Zamora, J. Señoret & J.C. Zamora, herb. Zamora 317; ibid., 24 Dec. 2010,
largest 63–80 µm diam), usually grouped to form 60–120 µm
B. Zamora, J.C. Zamora & J.C. Campos, herb. Zamora 470; ibid., 24 Dec.
2010, J.C. Zamora, B. Zamora & J.C. Campos, herb. Zamora 471; ibid., 6 Nov. diam rounded aggregates, intermixed with some 1.5–3.0 µm
2011, B. Zamora, herb. Zamora 498; ibid., 28 Dec. 2011, B. Zamora & J.C. wide, hyaline, branched, thin-walled, clamped hyphae. Pseudo
Zamora, herb. Zamora 500; ibid., 21 Oct. 2012, B. Zamora, J. Señoret & J.C. parenchymatous layer of thin-walled (≤ 1 µm thick), hyaline
to yellowish cells, variable in shape and size, about 20–60 × spicuous. Pseudoparenchymatous layer of thin-walled (mostly
15–40 µm. Fibrous layer with 2.0–4.5 µm wide, hyaline to very ≤ 1 µm thick) cells. Rhizomorphs with bipyramidal crystals of
pale yellowish, aseptate, straight or slightly sinuous, intertwined, COD, isolated or grouped in rose-like aggregates, and also
mostly unbranched, comparatively thick-walled (1.0 –1.5 µm some horn-like crystals of COM.
thick) hyphae, lumen visible. Mycelial layer double-layered; Ecology & Distribution — Only known from South America
inner layer consisting of 2.0 – 4.0 µm wide, strongly glued to- (Calonge et al. 2000, Cortez et al. 2008). It grows on humic
gether, more or less hyaline, branched, thin-walled and clamped soils of the ‘Tropical and subtropical moist broadleaf forests’
hyphae; outer layer with 1.5–2.5 µm wide, hyaline to somewhat biome in the Neotropic ecozone.
yellowish, aseptate, rarely branched, comparatively more or
Additional specimens examined. Argentina, Salta, La Candelaria, Palo
less thick-walled (0.5–1.5 µm thick) hyphae, lumen very narrow Quemado, Yungas, 12 Apr. 2012, L. Papinutti, G. Rolón & J.C. Zamora,
and difficult to perceive; myceliar projections indistinct from the MA-Fungi 86670. – Brazil, Rio Grande do Norte, Parque Estadual Dunas
outer myceliar layer. Rhizomorphs not studied. do Natal, sobre solo arenoso, 10 July 2004, I.G. Baseia, URFN-Fungos 229.
Ecology & Distribution — Known from only three specimens
collected in humid soils of the ‘Temperate broadleaf and mixed
Geastrum subsect. Geastrum
forests’ of the Paleartic ecozone.
Additional specimens examined (paratypes). UK (England), Hampshire, Key to species in Geastrum subsect. Geastrum
nr. New Milton, soil, Taxus, 2 Feb. 1995, D. & M. Nesbitt, K(M) 60288; Norfolk,
Surlingham, on soil, Quercus sp., June 2002, T.W. Dove, K(M) 99914.
1. Peristome clearly sulcate, with thick folds . . . . . . . . . . . . 2
1. Peristome fibrillose or transition between fibrillose and very
Geastrum leptospermum G.F. Atk. & Coker in Atkinson, Bot. finely sulcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Gaz. 36: 306. 1903, ‘Geaster leptospermus’ — Fig. 3, 9d 2. Basidiospores (4.8–)5.0–6.0(–6.5) µm, stalk mostly broader
Type. Lectotype (designated here, MBT198516): USA, North Carolina, than long, American species . . . . . . . . . . . . . . . G. smithii
Chapel Hill, on mosses covered living trunks of trees, Feb. 1903, W.C. 2. Basidiospores 5.5–7.0 µm, stalk slightly longer than broad,
Coker 2644, Atkinson Herbarium of Cornell University 14861b (slide) and Australasian species . . . . . . . . . . . . . . . . . . . G. aff. smithii
14861c (basidiomata), CUP! (since the collector number is the same in both
herbarium envelopes, we consider them as a single gathering). 3. Rhizomorph crystals mostly horn-like; mesoperidium as a
mealy cover made up mainly by generative hyphae, with
Synoptic description (based on Atkinson (1903), Coker & Couch scarce and small bipyramidal crystals (sometimes up to
(1928), Sunhede (1989), and the type specimens studied) — Exo- 75 µm diam, well-formed crystals, appear); stalk (1.0–)2.0–
peridium 8–12 mm diam in horizontal position, fornicate, with 5.0 mm high; peristome always fibrillose, indistinctly to more
(3–)4–7 non-hygrometric rays. Endoperidial body 2.5–5(–6.5) or less distinctly delimited; broadest capillitial hyphae (5.0–)
mm diam, pale to dark brown. Mesoperidium thin, with numerous 5.5–11.0(–15.0) µm wide . . . . . . . . . . . . . . . G. coronatum
but more or less sparse crystals. Largest mesoperidial crystals 3. Rhizomorph crystals mostly bipyramidal, grouped in rose-
of COD on the endoperidial surface 40–60 µm diam, bipyrami- like aggregates or not; mesoperidium formed by generative
dal. Peristome fibrillose, well-delimited. Stalk 0.2–0.5 mm high, hyphae and numerous bipyramidal crystals up to 127.5 µm
whitish. Basidiospores globose to slightly ovoid, 3.0–4.0 µm diam; stalk 1.0–2.5 mm high; peristome fibrillose to finely
diam, with (0.1–)0.2–0.3 µm high warts, ornamentation ver- sulcate, mostly distinctly delimited; broadest capillitial hy-
rucose. Pseudoparenchymatous layer of thin-walled (mostly phae (4.0 –)4.5–5.5 µm wide . . . . . . . . . G. thanatophilum
≤ 1 µm thick) cells. Rhizomorph crystals mainly oblique prisms
of COM, grouped in stellate aggregates; bipyramidal crystals
of COD also observed. Geastrum coronatum Pers., Syn. Meth. Fung.: 132. 1801
Ecology & Distribution — This rare species grows on mossy — Fig. 4, 10a
bark of living trees, and is only known from North Carolina Type. Lectotype (designated by Demoulin 1984): f. 2 of pl. XLVI in
(Atkinson 1903, Coker & Couch 1928), being found in the Schmidel (1793).
‘Temperate broadleaf and mixed forests’ biome of the Neartic
≡ Geastrum multifidum Pers., Neues Mag. Bot. 1: 86. 1794.
ecozone. ≡ Geastrum limbatum Fr., Syst. Mycol. 3: 15. 1829, ‘Geaster limbatus’,
Additional specimens examined. USA, North Carolina, Chapel Hill, on nom. illeg., Art. 52.1.
mosses covered living trunks of tree, woods, Feb. 1902 [probably an error for = Geastrum atratum F. Šmarda, Česká Mykol. 1: 74. 1947, ‘Geaster atra
1903], W.C. Coker 2319, Atkinson Herbarium of Cornell University 14861a, tus’. — Type: Unknown. In absence of original specimens, f. 2 in Šmarda (1947),
CUP (syntype). f. 25 of t. IX in Hollós (1904), or f. 44 in Lloyd (1902) can be selected as lecto-
type.
= Geastrum coronatum var. moelleri V.J. Staněk in Pilát, Flora ČSR B-1:
Geastrum ovalisporum Calonge & Mor.-Arr. in Calonge et al., 428, 789. 1958, ‘Mülleri ’ (the epithet ‘Mülleri ’ was corrected according to Art.
Bol. Soc. Micol. Madrid 25: 273. 2000 — Fig. 9e 60.6 and Rec. 60F). — Type: Hungary, Nagyerdö Csokás, Nagykörös, 14
Nov. 1955, I. Müller, holotype PRM 842868!
Type. Bolivia, Concepción, Piedra de Santa Teresita, en claros de bosque
tropical, sobre suelo arenoso con abundantes restos vegetales en descom- Synoptic description (based on Sunhede (1989) and studied spe-
posición, 5 Mar. 2000, B. Moreno-Arroyo & J. Gómez, holotype MA-Fungi cimens) — Exoperidium (33–)35–120(–135) mm diam in hori-
47184!
zontal position, arched, with (5–)7–12(–15) sometimes falsely
Synoptic description (based on Calonge et al. (2000) and stud- hygrometric rays. Endoperidial body 9–43 mm diam, brownish
ied specimens) — Exoperidium 26 – 45 mm diam in horizontal to blackish. Mesoperidium as a dense mealy cover, mostly with
position, arched, with 5 – 8 non-hygrometric rays. Endoperidial scarce or indistinct crystalline matter, but sometimes with well-
body 8 –15 mm diam, brown to almost black. Mesoperidium developed crystals. Largest mesoperidial crystals of COD on
thin, with rather sparse crystals. Largest mesoperidial crys the endoperidial surface 2–53(–75) µm diam, bipyramidal. Peri
tals of COD on the endoperidial surface 37.5 –78 µm diam, stome fibrillose, distinctly or indistinctly delimited. Stalk stout,
bipyramidal (bipyramids and bipyramidal prisms), sometimes (1.0–)2.0–4.5(–5.5) mm high, brownish or greyish to almost
irregular. Peristome fibrillose, well-delimited. Stalk 1.3 – 2.0 black, rarely cream coloured. Basidiospores globose, (5.0–)5.5–
mm high, whitish. Basidiospores ovoid, 3.0 – 4.0 × 2.5 –3.5 µm, 6.5 (–7.0) µm diam, with 0.5 –1.0 µm high warts, orna-
with 0.1– 0.3 µm high warts, ornamentation verrucose, incon- mentation verrucose. Broadest capillitial hyphae (5.0 –)5.5 –
11.0(–15.0) µm wide. Pseudoparenchymatous layer of thin- MA-Fungi 83783; Córdoba, Punilla, Hayke, a 3 km de Icho Cruz, por ruta
walled (mostly ≤ 1 µm thick) cells. Rhizomorphs mainly with provincial N°14, sobre mantillo bajo espinillo, 20 Jan. 1990, L. de Toledo
843, CORD 843; La Rioja, Sañogasta, S29°18'51.43" W67°35'37.36", on
thin horn-like crystals of COM, forming arachnoid structures.
abundant leaf litter of diverse herbaceous plant species, 27 Mar. 2008,
Ecology & Distribution — Confirmed records are known
L. Papinutti & G. Rolón, BAFC 51945. – USA, New Mexico, 6 mi. W of Corona,
from calcareous and siliceous soils, mostly on humic places, 4 Sept. 1941, Long & Stouffer 9505, LPS 29986; ibid., 15 Sept. 1941, Long
and from several biomes (‘Boreal forests/taiga’, ‘Temperate & Stouffer 9662, LPS 29801; New Mexico, 2 mi. NE of Corona, Long &
conifer forests’, ‘Temperate broadleaf and mixed forests’, and Stouffer, 17 Apr. 1942, LPS 29979; ibid., 14 Jan. 1942, Long & Stouffer
‘Mediterranean forests, woodlands and scrub’) of the Neartic 9993, LPS 30189.
and Paleartic ecozones (Sunhede 1989, Calonge 1998, Bates
2004). Also reported from Paleotropical (Bottomley 1948) and Geastrum aff. smithii Lloyd — Fig. 10d
Australasian (Cunningham 1944) ecozones.
Synoptic description — Exoperidium 42–47 mm diam in hori
Additional specimens examined. Czech Republic, Ýmel, 1972, E. Futó,
MICH 72513. – Spain, Madrid, Casa de Campo, enfrente del Club de Campo,
zontal position, arched, with 8–9 not hygrometric or falsely hygro-
bajo Pinus sp., 12 Dec. 2010, L. Penelas, herb. Zamora 464; Madrid, Villavi- metric rays. Endoperidial body 13–15 mm diam, dark brownish
ciosa de Odón, parque cercano al Castillo, sandy, siliceous, and humic soil, grey. Mesoperidium well-developed, with abundant crystals.
under Ulmus minor and Cupressus arizonica, 24 Dec. 2003, J.C. Zamora, Largest mesoperidial crystals of COD on the endoperidial sur-
herb. Zamora 37; ibid., under Ulmus minor, 19 Sept. 2006, B. Zamora & face 120–160 µm diam, bipyramidal. Peristome sulcate, sharply
J.C. Zamora, herb. Zamora 158; ibid., under Quercus robur, 28 Oct. 2006, delimited, flat to very broadly conical, with 25–29 folds. Stalk
J.C. Zamora, herb. Zamora 181; ibid., under Cupressus sempervirens, Ul
stout, 2.0 – 2.5 mm high, light to dark brown. Basidiospores
mus minor, Populus sp., Calocedrus sp., and Celtis australis, siliceous soil,
25 Nov. 2007, B. Zamora & J.C. Zamora, herb. Zamora 266; ibid., under globose, 5.5–7.0 µm diam, with 0.5–0.8 µm high warts, ver-
Cupressus sp., Cedrus sp., Ulmus minor, and Populus sp., 27 Dec. 2009, rucose ornamentation. Broadest capillitial hyphae 5.0 – 5.5
S. Pardillo, P.L. Aznar & J.C. Zamora, herb. Zamora 364; ibid., under Quercus µm diam. Pseudoparenchymatous layer of thin-walled (mostly
robur, 24 Dec. 2010, B. Zamora, J.C. Zamora & J.C. Campos, herb. Zamora ≤ 1 µm thick) cells.
476; ibid., under Ulmus minor and Pinus sp., 7 May 2011, B. Zamora, J. Se- Ecology & Distribution — Insufficiently known; a single her-
ñoret & J.C. Zamora, herb. Zamora 484; ibid., under Ulmus minor and Cu-
barium collection from the ‘Temperate grasslands, savannas
pressus sempervirens, 13 Nov. 2011, B. Zamora & J.C. Zamora, herb. Zamora
549; ibid., under Cupressus sp. and Ulmus minor, 13 Nov. 2011, B. Zamora
and shrublands’ biome of the Australasian ecozone has been
& J.C. Zamora, herb. Zamora 576. – Sweden, Gotland, Bunge parish, Bunn, studied.
dead anthill, 5 Dec. 1970, S. Sunhede, herb. Sunhede 7601; Gotland, Gothem
Additional specimens examined. Australia, New South Wales, Riverina,
parish, Jusarve skog, on an anthill, mixed forest with Pinus sylvestris, Picea
Ringwood State Forest, SANDS Plot 26, 11 June 2001, J. Trappe 26384A,
abies, and Quercus robur, on calcareous soil, 30 Sept. 2011, J.C. Zamora,
CANB 748746 (duplo in OSC).
herb. Zamora 522. – USA, Arizona, Coconino Co., Walnut Canyon National
Monument, Pinus edulis and Juniperus, 3 May 1995, J. States AEF 1443,
MICH 28567. Geastrum thanatophilum J.C. Zamora, sp. nov. — MycoBank
Nomenclatural notes — Demoulin (1984) made homotypic MB810501; Fig. 4, 10b
the names G. coronatum and G. multifidum by selecting one of Type. USA, Wisconsin, Dane Co., Mazomanie cemetery, between old
the figures cited under ‘G. multifidum var. β’ in Persoon (1794) as juniper and a headstone, 29 Sept. 2007, J.J. Steinke & P. Watson, holotype
type for both, while, according to Art. 32.1(b), the name ‘G. multi- MICH 72012!
fidum var. β’ itself cannot be considered as validly published in Etymology. From the Greek words Θάνατος (transliterated ‘Thanatos’, god
the infraspecific rank, see also Ex. 4 of Art. 9.5. of death) and φίλος (transliterated ‘philos’, friend), because the two studied
collections came both from cemeteries.
Geastrum smithii Lloyd, Bull. Lloyd Libr. Bot. 5: 21. 1902, Diagnosis — Exoperidium 22–46 mm diam in horizontal posi-
‘Geaster’ — Fig. 10c tion, arched, with 6–11(–14) sometimes falsely hygrometric rays.
Type. Lectotype (designated here, MBT198517): USA, Florida, New Endoperidial body 7–16 mm diam, greyish cream to brownish
Smyrna, Mrs. Sams, Smithsonian Institution (Herb. Lloyd) 22731, BPI 705991 grey. Mesoperidium as a dense mealy cover with big crystals.
(photo!). Largest mesoperidial crystals of COD on the endoperidial sur-
face 55–127.5 µm diam, bipyramidal. Peristome fibrillose to
Synoptic description (based on Bates (2004) and studied spe
obscurely sulcate, then with up to 35 folds, mostly well-delim-
cimens) — Exoperidium (14 –)30 – 57 mm diam in horizontal
ited. Stalk stout, 1.0–2.5 mm high, normally greyish to rather
position, arched, with 7–12 sometimes falsely hygrometric
rays. Endoperidial body 7– 23 mm diam, pale brown to black- dark brown. Basidiospores globose, (4.5–)5.0–6.0(–6.5) µm
ish. Mesoperidium well-developed, with abundant big and small diam, with (0.3–)0.4–0.8(–1.0) µm high warts, ornamentation
crystals. Largest mesoperidial crystals of COD on the endo verrucose. Broadest capillitial hyphae (4.0–)4.5–5.5 µm wide.
peridial surface about (20–)50–100 µm diam, bipyramidal. Peri Pseudoparenchymatous layer of thin-walled (mostly ≤ 1.2 µm
stome sulcate, sharply delimited, normally flat to very broadly thick) cells. Rhizomorphs with bipyramidal crystals of COD,
conical, with 16 – 30 folds. Stalk stout, 1.5 – 3.0 mm high, light isolated or grouped in rose-like aggregates.
to dark brown. Basidiospores globose, (4.8 –)5.0 – 6.0(– 6.5) Macroscopic characteristics — Unexpanded basidiomata
µm diam, with (0.4 –)0.5 – 0.7(– 0.8) µm high warts, verrucose 15–26 mm diam, subglobose, with a rounded apex or a flat
ornamentation. Broadest capillitial hyphae 5.0 – 6.5 µm diam. umbo. Exoperidium splitting in 6–11(–14) subequal to unequal
Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm rays, 9–41 mm diam apparently, 22–46 mm diam when forced
thick) cells. in horizontal position, arched, not truly hygrometric but young
Ecology & Distribution — Known from ‘Temperate conifer rays may bent towards the endoperidial body when the myce-
forests’, ‘Tropical and subtropical grasslands, savannas and lial layer peels off. Mycelial layer thick, whitish to pale cream,
shrublands’, and ‘Deserts and xeric shrublands’ biomes, of strongly intermixed with debris from the substrate, more or less
the Neartic and Neotropic ecozones (Lloyd 1902, Bates 2004). strongly adhered to the fibrous layer but peeling-off in some
Records from the Australasian ecozone (Cunningham 1944) parts. Fibrous layer more or less papyraceous to coriaceous
probably belong to a different taxon (see below). when denuded, whitish to pale-cream coloured. Pseudo
Additional specimens examined. Argentina, Córdoba, Cruz del Eje, Villa parenchymatous layer pale cream to greyish cream, mostly
de Soto, Cupressus sp., 6 Apr. 2012, L. Papinutti, J.C. Zamora & G. Rolón, uncracked, < 0.5 mm thick in dry state, about 1.5 – 2.5 mm
thick when hydrated, not persisting in old basidiomata. Endo Peristome fibrillose to somewhat sulcate, with 0–35 obscure
peridial body globose to subglobose, rarely irregular, mostly folds, concolorous with the endoperidial surface, broadly coni-
wider than high, 7–16 mm diam, greyish cream to brownish cal to almost flat, up to 1 mm high, mostly distinctly delimited
grey; endoperidial surface glabrous or almost so, in newly ex- but specimens with indistinctly delimited peristomes occur.
panded basidiomata covered with abundant collapsed hyphae Stalk present, comparatively stout, 1.0–2.5 mm high, greyish
and some big, bipyramidal crystals, gradually disappearing. to rather dark brown, very rarely pale coloured. Apophysis ab-
Fig. 10   Morphological characters in Geastrum subsect. Geastrum and Geastrum subsect. Quadrifida pro parte (G. kuharii). a. G. coronatum (Zamora 266);
b. G. thanatophilum (holotype); c. G. smithii (MA-Fungi 83783); d. G. aff. smithii (CANB 748746); e. G. kuharii (holotype). — a1, b1, c1, d1, e1. Basidiomata
habit; a2, b2, c2, d2, e2. detail of the peristome; a3, b3, c3, d3, e3. detail of the stalk and apophysis; a4, b4, c4, d4, e4. mesoperidial COD crystals on the
endoperidial surface, in a4 almost no crystalline matter present; a5, b5, c5, d5, e5. basidiospores. — Scale bars: a1, b1, c1, d1, e1 = 5 mm; a2, b2, c2, d2,
e2, a3, b3, c3, d3, e3 = 2 mm; a4, b4, c4, d4, e4 = 10 µm; a5, b5, c5, d5, e5 = 2 µm.
sent or poorly developed, concolorous with the endoperidium. Synoptic description (based on Hollós (1904), Staněk (1958),
Columella intruding about 1/3–1/2 into the glebal mass. Mature Sunhede (1989), and studied specimens) — Exoperidium
gleba dark greyish brown. 10 – 25 mm diam in horizontal position, saccate to arched
Microscopic characteristics — Basidia narrowly ellipsoid, when hydrated, with 5 –12 strongly hygrometric rays. Endo
more or less lageniform to more or less lecythiform, (12.5–)14– peridial body 2 – 8 mm diam, whitish to pale greyish brown.
20 × 5.0 –7.5 µm, with 4 – 8 short sterigmata. Basidiospores Mesoperidium as a thin layer of pruina. Largest mesoperidial
globose to subglobose, (4.5–)5.0–6.0(–6.5) µm diam, brownish crystalline aggregates of COM on the endoperidial surface
to yellowish brown, with (0.3 –)0.4 – 0.8(–1.0) µm high brown about 25 – 30 µm diam, bipyramidal crystals of COD scarce.
warts, ornamentation verrucose. Broadest capillitial hyphae Peristome fibrillose, distinctly delimited. Stalk absent or indis-
(4.0–)4.5 – 5.5 µm wide, aseptate, very rarely branched, nor- tinct. Basidiospores globose to subglobose, 5.0 – 6.0 µm diam,
mally straight, thick-walled (2 – 2.5 µm thick), with narrow lu- with 0.4 – 0.7 µm high warts, ornamentation verrucose. Pseu
men, more or less visible; tips acute to rounded; surface often doparenchymatous layer very persistent, with clearly thick-
covered with debris. Endoperidium composed of 2.0 –7.0 µm walled (> 1.5 µm thick) cells.
wide, yellowish to pale brownish, aseptate, mostly unbranched, Ecology & Distribution — This species mainly grows in open
slightly sinuous, strongly intertwined, thick-walled hyphae, with places with scarce trees (Staněk 1958, Sunhede 1989), and
lumen mostly visible; protruding hyphae not seen. Peristomal most records are known from the ‘Temperate broadleaf and
hyphae 2.5 – 8.0 µm wide, yellowish brown to brownish, asep- mixed forests’ biome of the Paleartic ecozone. In addition,
tate, mostly unbranched, thick-walled (1.0–3.0 µm thick), lumen we have studied one sample from the ‘Mediterranean forests,
often visible, more or less straight to rather sinuous, narrowing woodlands and scrub’ biome of the Paleartic ecozone.
at base and apex, tips mostly rounded, a few acute. Mesoperi
Additional specimens examined. Czech Republic, Reporyje, u Prahy, slope
dium present on the endoperidium and pseudoparenchymatous with Stipa capillata and Gastrosporium simplex, 8 May 1955, Kotlaba, Pouzar
layer surfaces, consisting of rather large, bipyramidal crystals & Staněk, herb. Sunhede 5993. – Spain, Toledo, Argés, en claro arenoso de
of COD, the largest 55 –127.5 µm diam, intermixed with abun- encinar, 8 Dec. 2008, J. De Esteban, herb. Zamora 611.
dant 1.5–4.0 µm wide, hyaline, branched, thin-walled, clamped
Nomenclatural notes — Although Sunhede (1989) desig-
hyphae. Pseudoparenchymatous layer of thin- to slightly thick-
nated a neotype (BP 23244) for this species, original material
walled (0.5 –1.2 (–1.5) µm thick), hyaline to yellowish cells,
from herb. Hollós, collected before publication of the name of
variable in shape and size, about 15 –112 × 11– 50 µm diam.
the species, exists in Stockholm (S F16379). Therefore, the
Fibrous layer with 1.5 – 4.5(– 5.0) µm wide, hyaline to very pale
neotype designated by Sunhede (1989) is superseded follow-
yellowish, aseptate, straight or slightly sinuous, intertwined,
ing Art. 9.19(a), and the collection S F16379 is designated as
mostly unbranched, comparatively thick-walled (0.5 –2.0 µm
lectotype.
thick) hyphae, lumen more or less visible. Mycelial layer double-
layered; inner layer consisting of 2.0 – 3.0 µm wide, strongly
glued together, more or less hyaline, branched, thin-walled and Geastrum subsect. Quadrifida J.C. Zamora in Zamora et al.,
clamped hyphae; outer layer with 1.5 – 4.0 µm wide, hyaline to Taxon 63, 3: 491. 2014
somewhat yellowish, aseptate, rarely branched, comparatively
more or less thick-walled (0.7– 2.0 µm thick) hyphae, lumen Key to species in Geastrum subsect. Quadrifida
often very narrow and difficult to observe. Rhizomorphs with
bipyramidal crystals of COD, isolated or grouped in rose-like 1. Exoperidium fornicate, with (3 –)4 – 5(– 6) rays, margin of
aggregates. rays conspicuously rolled out . . . . . . . . . . . G. quadrifidum
Ecology & Distribution — The two studied collections have 1. Exoperidium not fornicate, with normally more than 4 rays,
been found growing on coniferous litter of anthropized envi- margin of rays from rolled in to slightly rolled out . . . . . . . 2
ronments (cemeteries), being difficult to known the ecological 2. Basidiomata rather big and stout (exoperidium 45–62 mm
requirements of the species. Geographically, those places are diam and endoperidial body 13–21 mm diam); mesoperidium
located in the ‘Temperate broadleaf and mixed forests’ biome with both abundant generative hyphae and big crystals or
of the Neartic ecozone. crystalline aggregates . . . . . . . . . . . . . . . . . . . . G. kuharii
Additional specimens examined (paratypes). USA, Wisconsin, Crawford 2. Basidiomata smaller and slender (exoperidium (9–)11–45
Co., Prairie du Chien, Prairie du Chien Cemetery, in duff between mature ( – 60) mm diam and endoperidial body 3 –14(–15.5) mm
juniper and headstone, 14 Jan. 2006, J.J. Steinke, MICH 72014. diam); mesoperidium with few generative hyphae and abun-
dant or scarce crystals or crystalline aggregates . . . . . . . 3
Geastrum subsect. Hungarica J.C. Zamora, subsect. nov. — 3. Endoperidial surface with a rather sparse mesoperidial crys-
MycoBank MB810518 tal cover; largest mesoperidial crystals or crystalline aggre-
gates < 70(–95) µm; stalk mostly dark coloured . . . . . . . .
Type. Geastrum hungaricum Hollós.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. marginatum
Diagnosis — Exoperidium more or less saccate to arched when 3. Endoperidial surface with a dense mesoperidial crystal
hydrated, strongly hygrometric (pseudoparenchymatous layer cover; largest mesoperidial crystals or crystalline aggregates
with thick-walled cells). Mycelial layer encrusting debris, double- > (65–)70 µm; stalk light or dark coloured . . . . . . . . . . . . 4
layered. Endoperidial body sessile. Endoperidial surface gla- 4. Apophysis very well-developed, almost ring-like; stalk 1.5–
brous and covered by a well-developed mesoperidium, com- 2.3 mm high . . . . . . . . . . . . . . . . . . . . . (G. calceum s.l.) 5
posed by generative hyphae and abundant crystalline ag- 4. Apophysis absent to more or less conspicuous, but not ring-
gregates of COM, with scarce bipyramidal crystals of COD. like; stalk 0.5–2.0 mm high . . . . . . . . . . . . . . . . . . . . . . . 6
Peristome fibrillose and distinctly delimited.
5. Stalk slender, 2.0 – 2.3 mm high, cream coloured to pale
brown; basidiospores 4.5–6.0 µm, ornamentation 0.5–0.8
Geastrum hungaricum Hollós, Math. Termeszettud. Ertes. 19: µm high . . . . . . . . . . . . . . . . . . . . . . . . . . G. cf. calceum1
507. 1901, ‘Geaster hungaricus’ — Fig. 9f 5. Stalk more or less slender or not, 1.5–2.0 mm high, brown-
Type. Lectotype (designated here, MBT198541): Hungary, Nagy-Körös, ish; basidiospores 5.5–6.5 µm, ornamentation (0.5–)0.6–
3 Dec. 1898, in arenos, Ex Herbario Dr. L. Hollós, Kecskemét, S F16379! 1.0(–1.2) µm high . . . . . . . . . . . . . . . . . . .G. cf. calceum 2
Fig. 11   Morphological characters in Geastrum subsect. Quadrifida (except G. kuharii). a. G. austrominimum (holotype); b. G. cf. calceum1 (MA-Fungi 83761);
c. G. cf. calceum2 (UFRN-Fungos 723); d. G. granulosum (Sunhede 7746); e. G. marginatum (MA-Fungi 86669); f. G. quadrifidum (Zamora 590 and MA-Fungi
86671). — a1, b1, c1, d1, e1, f1. Basidiomata habit; a2, b2, c2, d2, e2, f2. detail of the peristome; a3, b3, c3, d3, e3, f3. detail of the stalk and apophysis;
a4, b4, c4, d4, e4, f4. mesoperidial crystalline matter on the endoperidial surface: a4 shows crystalline aggregates of COM scales, b4, c4, d4, e4, f4 show
bipyramidal crystals of COD; a5, b5, c5, d5, e5, f5. basidiospores. — Scale bars: a1, b1, c1, d1, e1, f1 = 5 mm; a2, b2, c2, d2, e2, f2, a3, b3, c3, d3, e3, f3 =
2 mm; a4, b4, c4, d4, e4, f4 = 10 µm; a5, b5, c5, d5, e5, f5 = 2 µm.
6. Stalk more or less dark coloured, pale brown to rather dark without debris. Endoperidium composed of 2.0–5.5 µm wide,
brownish grey; mesoperidium normally with abundant and pale yellowish to yellowish brown, aseptate, mostly unbranched,
rather big, rounded, yellowish aggregates of COM; some slightly sinuous, strongly intertwined, thick-walled hyphae, lu-
bipyramidal COD crystals often present, but being rarely men mostly visible; protruding hyphae not seen. Peristomal hy
the main type; Australasian species . . . G. austrominimum phae 4.5–10(–15) µm wide, light brown, aseptate, unbranched
6. Stalk light-coloured, whitish to cream coloured; mesoperi except for scarce short branches mostly near the base, thick-
dium mostly with big bipyramidal crystals of COD; crystalline walled (1.5–4.0 µm thick), lumen often visible, often sinuous
aggregates of COM also often present, but normally much to wavy, narrowing at base and apex, tips mostly rounded, a
less abundant or inconspicuous; Holartic species . . . . . . . few acute. Mesoperidium present on the endoperidium and
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. granulosum pseudoparenchymatous layer surfaces, composed by rather
large, rounded, crystalline aggregates of COM scales, the
Geastrum austrominimum J.C. Zamora, sp. nov. — Myco- largest 30–105 µm diam, less abundant bipyramidal crystals
Bank MB810502; Fig. 6, 11a of COD, the largest 20–130(–200) µm diam, rarely dominant,
and sparse 1.5–3.0 µm wide, hyaline, branched, thin-walled,
Type. Australia, New South Wales, Riverina, Savernake Station, SANDS
clamped hyphae. Pseudoparenchymatous layer of thin-walled
Plot 8, S35°44' E146°03', 11 June 2001, A. Sloan, S. Reilly, J. Trappe & A.
Giachini, Trappe 26378, holotype CANB 748741!, isotype in OSC.
(≤ 1 µm thick), hyaline to pale yellowish cells, variable in shape
and size, about 11–70 × 10–40 µm. Fibrous layer with 3.0–8.0
Etymology. The epithet is a combination of the prefix ‘austro-’ and the µm wide, mostly yellowish, aseptate, straight or slightly sinuous,
ending ‘-minimum’, meaning that this species corresponds to the Australasian
intertwined, mostly unbranched, thick-walled (1.5–3.0 µm thick)
specimens for which the name G. minimum has been used.
hyphae, lumen visible. Mycelial layer double-layered; inner layer
Diagnosis — Exoperidium 17– 35 mm diam in horizontal posi- consisting of 1.5–3.5 µm wide, strongly glued together, more
tion, arched, with 6 –11(–13) not hygrometric rays. Endoperi or less hyaline, branched, thin-walled and clamped hyphae;
dial body 5 –10 mm diam, greyish cream to brownish grey. outer layer with 1.0–4.0 µm wide, hyaline to somewhat yel-
Mesoperidium as a dense layer of crystalline matter. Largest lowish, aseptate, rarely branched, comparatively more or less
mesoperidial crystalline aggregates of COM scales 30 –105 thick-walled (0.5–2.0 µm thick) hyphae, lumen very narrow and
µm diam, often very abundant and densely covering the endo- difficult to perceive. Rhizomorphs with rose-like aggregates of
peridial surface. Largest mesoperidial crystals of COD on the bipyramidal crystals of COD.
endoperidial surface 20 –130(– 200) µm diam, bipyramidal, Ecology & Distribution — The studied specimens came from
mostly sparse, rarely dominant. Peristome fibrillose, flat to ‘Temperate grasslands, savannas and shrublands’ and ‘Tem-
broadly conical, mostly well-delimited. Stalk more or less stout, perate broadleaf and mixed forests’ biomes of the Australasian
0.5–2.0 mm high, normally brownish (paler when old). Basidio ecozone.
spores globose, 4.5–6.5 µm diam, with 0.4–1.0 µm high warts,
Additional specimens examined (paratypes). Australia, New South Wales,
ornamentation verrucose to irregularly pilate. Broadest capillitial South Western Plains, Riverina, Savernake Station, SANDS Plot 8, S35°45'
hyphae 5.0–8.5 µm wide. Pseudoparenchymatous layer of thin- E46°01', under Eucalyptus microcarpa, 7 Nov. 2000, J. Trappe 26610, MEL
walled (mostly ≤ 1 µm thick) cells. Rhizomorphs with rose-like 2276089; Victoria, Neds Corner Station, NW corner of property, VicMuseum
aggregates of bipyramidal crystals of COD. pitfall line 004, S34.1137° E141.211°, woodland, 25 Nov. 2011, T. Lebel 2433,
MEL 2358014; Victoria, Midlands, Joel Bushland Reserve on south side of
Macroscopic characteristics — Unexpanded basidiomata not Stawell-Landsborough Rd, S37°02' E142°55', Vic. Grid Ref. J 3, 27 July
seen. Exoperidium divided in 6 –11(–13) more or less equal to 1996, I.R. McCann, MEL 2103688; Victoria, Murray Mallee, Hattah-Kulkyne
unequal rays, 11–32 mm diam apparently, 17–35 mm diam when National Park, off Mournpall Track, S34°44' E142°21', Vic. Grid Ref. A 45, on
forced in horizontal position, arched, not hygrometric. Mycelial dune under Callitris, 7 June 1998, I.R. McCann & T.G. Argall, MEL 2104353;
Victoria, Midlands, Kamarooka State Park, East Kamarooka Rd, 500 m
layer thin to more or less thick, whitish to pale cream, strongly
N of Boobialla Track, E269185 N5953222, Claridge site 231 - (no EVC),
intermixed with debris from the substrate, strongly adhered to S36°32'18" E144°25'18", 180 m, Vic. Grid Ref. M 30, flat, gravelly-sediment
the fibrous layer but peeling-off in certain parts. Fibrous layer pa- mallee, Eucalyptus behriana, E. froggattii, 4 Aug. 2002, J. Trappe 27826,
pyraceous when denuded, whitish to cream coloured. Pseudo- MEL 2271749; Victoria, Midlands, Black Range, S of Stawell, in Bunjils Cave
parenchymatous layer pale cream to brownish, sometimes Reserve, S37°08'13" E142°43'59", Vic. Grid Ref. J 2, near long-leaf Box,
superficially cracked, < 0.5 mm thick in dry state, not persisting 1 Apr. 2001, I.R. McCann & T.G. Argall, MEL 2292062; Victoria, Neds Corner
in old basidiomata. Endoperidial body globose to subglobose, Station, near Snake Lagoon track, S34°08'44.3" E141°23'51.3", blackbox
woodland, 30 Nov. 2011, T. Lebel 2372 & A. Pay, MEL 2358047.
rarely irregular, 5 –10 mm diam, pale cream to brownish; endo
peridial surface glabrous or almost so. Mesoperidium mainly
composed of rather dense and persistent crystalline matter, Geastrum calceum Lloyd, Mycol. Writings 2: 311. 1907,
often with abundant yellowish, rounded crystal aggregates, ‘Geaster calceus’ s.l. — Fig. 6
intermixed with some whitish crystals that are rarely dominant. Type. South Africa, O. Pazschke, C.G. Lloyd Mycological Collection,
Peristome fibrillose, of the same colour as the endoperidial Smithsonian Institution 57280, BPI 704885 (photo!, probably a syntype).
surface, mostly flat to broadly conical and distinctly delimited.
Stalk present, not particularly slender, 0.5–2 mm high, normally Synoptic description of G. cf. calceum1 (Fig. 11b) — Exoperi
pale brown to rather dark brownish grey, but may be paler in old dium 41–45 mm diam in horizontal position, arched, with 8–9
basidiomata. Apophysis present or absent, the same colour as not hygrometric rays. Endoperidial body 11–14 mm diam, pale
the endoperidium or slightly darker. Columella intruding about brown to brown. Apophysis almost ring-like. Mesoperidium as
1/3–1/2 into the glebal mass, with a more or less rounded apex. a more or less dense layer of crystals. Largest mesoperidial
Mature gleba dark greyish brown. crystals of COD on the endoperidial surface 92–115 µm diam,
Microscopic characteristics — Basidia not seen. Basidio bipyramidal. Largest mesoperidial crystalline aggregates of
spores globose to subglobose, 4.5 – 6.5 µm diam, brownish to COM scales 13 – 30 µm diam, often very inconspicuous or
yellowish brown, with 0.4–1.0 µm high brown warts, ornamenta- doubtful. Peristome fibrillose, broadly conical to almost flat,
tion verrucose to irregularly pilate. Broadest capillitial hyphae well-delimited. Stalk more or less slender or not, 1.5–2.0 mm
5.0–8.5 µm wide, aseptate, normally not branched and more high, brownish. Basidiospores globose, 5.5–6.5 µm diam, with
or less straight, thick-walled (2.5 – 3.5 µm thick), with narrow (0.5–)0.6–1.0(–1.2) µm high warts, ornamentation verrucose
but mostly visible lumen; tips acute to rounded; surface with or to irregularly pilate. Broadest capillitial hyphae 5.5 – 6.0 µm
wide. Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 forests’, and ‘Mediterranean forests, woodlands and scrub’) of
µm thick) cells. the Neartic and Paleartic ecozones.
Ecology & Distribution — Insufficiently known; the single Additional specimens examined. Germany, Oestrich, Nassau, c. Buden-
studied herbarium collection came from an anthropized environ- heim, in pinetis, Fuckel, herbier Barbey-Boissier 2138, UPS; c. Budenheim,
ment, geographically located in the ‘Tropical and subtropical in pinetis, 1265, UPS F-127433. – Spain, Madrid, Alcalá de Henares, 10 Dec.
grasslands, savannas and shrublands’ biome of the Neotropic 1993, G. Manteiga, MA-Fungi 32165; Madrid, Alcalá de Henares, cerro del
ecozone. Viso, bajo Pinus halepensis, terreno básico, 12 Nov. 2011, R. Suárez Rico,
herb. Zamora 600; Madrid, Alcalá de Henares, bajo Pinus halepensis, suelo
Additional specimens examined. Argentina, Tucumán, San Miguel de calizo superficialmente quemado, 20 Nov. 2011, J. Hernanz, B. Zamora &
Tucumán, parque Lillo, Nov. 2011, E. Grassi, MA-Fungi 83761. J.C. Zamora, herb. Zamora 550; Madrid, Arganda del Rey, dehesa ‘El Car-
rascal’, 30 Nov. 2003, L. Rubio, MA-Fungi 59067; ibid., bajo conífera inde-
Synoptic description of G. cf. calceum2 (Fig. 11c) — Exoperi terminada, 18 Nov. 2006, L. Penelas, herb. Zamora 191; Madrid, Arganda
dium 38–40 mm diam in horizontal position, arched, with 7–8 del Rey, bajo Quercus coccifera, 30 Nov. 2005, L. Rubio, MA-Fungi 69175;
not hygrometric rays. Endoperidial body 10–12 mm diam, grey- Madrid, Cerro de los Ángeles, en pinar de Pinus halepensis, 8 Dec. 1991,
ish cream to more or less dark brown. Apophysis almost ring- A. Guerra, MA-Fungi 28118; Murcia, Fuente del Obispo, sierra de la Fuensan-
like. Mesoperidium as a dense layer of crystals. Largest meso ta (sierra Carrascoy), 30SXG6497, 500 m, entre agujas de Pinus halepensis,
peridial crystals of COD on the endoperidial surface 80–85 µm 9 Nov. 1978, M. Honrubia, MA-Fungi 42282 (ex MUB 1653); ibid., pinar de
Pinus halepensis, en fisuras de rocas, 1 Mar. 1980, X. Llimona, MHG 2975,
diam, bipyramidal. Largest mesoperidial crystalline aggregates MA-Fungi 42283 (ex MUB 1656); ibid., pinar aclarado de Pinus halepensis,
of COM scales 12–15 µm diam, often very inconspicuous or 14 Nov. 1978, M. Honrubia, MHG 343, MA-Fungi 42284 (ex MUB 1654);
doubtful. Peristome fibrillose, broadly conical to almost flat, ibid., pinar de Pinus halepensis, 22 May 1978, M. Honrubia, MHG 311, MA-
well-delimited. Stalk slender, 2.0–2.3 mm high, cream coloured Fungi 42287 (ex MUB 1658); ibid., pinar de Pinus halepensis, 14 Nov. 1978,
to pale brown. Basidiospores globose, 4.5–6.0 µm diam, with M. Honrubia, MHG 343, MA-Fungi 43290 (ex MUB 343); Soria, cañón del
0.5–0.8 µm high warts, ornamentation verrucose to irregularly río Lobos, bajo Pinus sp. y Juniperus thurifera, 9 Nov. 1992, T. Almaraz,
MA-Fungi 32393; Tarragona, La Cenia, 10 Nov. 1973, J.M. Alemany, MA-
pilate. Broadest capillitial hyphae 7.0–8.0 µm wide. Pseudo
Fungi 42280 (ex MUB 1650); Teruel, puerto de Alcalá de la Selva, 28 Oct.
parenchymatous layer of thin-walled (mostly ≤ 1 µm thick) cells. 1993, F.D. Calonge, MA-Fungi 32164; Teruel, Bezas, campo descubierto, 27
Ecology & Distribution — Insufficiently known; studied speci Oct. 1993, F.D. Calonge, MA-Fungi 32163; Teruel, Formiche, 24 Oct. 1990,
mens came from ‘Tropical and subtropical moist broadleaf F.D. Calonge, MA-Fungi 31162; Teruel, Linares de Mora, 25 Oct. 1995, bajo
forests’ biome of the Neotropic ecozone. Pinus sylvestris, F.D. Calonge, MA-Fungi 34054; Teruel, Valdelinares, bajo
Pinus sylvestris, 28 Oct. 1994, F.D. Calonge, MA-Fungi 33261; Valladolid,
Additional specimens examined. Brazil, Rio Grande do Norte, Natal, Portillo, bajo Pinus pinea, 20 Dec. 1999, A. García, MA-Fungi 47907. –
Tricha da Geologia, P. Dunas, 14 Apr. 2010, M.M.B. Barbosa, B. Tomio, T. Sweden, Öland, Gårdby parish, N. Gårdby, open sandy grazed and trampled
Lockwood & I.G. Baseia, UFRN-Fungos 723. – Costa Rica, Guanacaste, ground, with i.a. Veronica spicata, near forest border (Pinus sylvestris),
Parque Nacional Santa Rosa, Bosque de San Emilio, 300 m, 15 Apr. 2003, 28 Aug. 2012, J.C. Zamora & S.I. Sunhede, herb. Sunhede 7746. – USA,
M. Mata 1279, MA-Fungi 65435 (duplo INB 3721248). Arizona, Coconino Co., Walnut Canyon National Monument, Pinus edulis,
14 Oct. 1986, J. States AEF 810, MICH 28210; ibid., in open area around
Additional remarks — Although requested on loan, type ma debris, Pinus edulis and Juniperus, 17 Oct. 1986, J. States AEF 518, MICH
terial of G. calceum was not available for detailed morphological 28119a; Wisconsin, Waukesha Co., NW corner of Hwy E and Hwy 99, very
study, which prevented us to make a proper synoptic description stony, specimens in needle duff around two juniper trees near top of hill, 25
because of the absence of important morphological characteris Oct. 1998, J.J. Steinke, MICH 72010.
tics, such as micromorphological data, in the protologue.
Nomenclatural notes — At UPS herbarium there are three
herbarium specimens (UPS F-127433 (1265), UPS F-127434
Geastrum granulosum Fuckel, Jahrb. Nassauischen Vereins (Fuckel 317), and Herbier Barbey-Boissier 2138) collected by
Naturk. 15: 41. 1860, ‘Geaster granulosus’ — Fig. 6, 11d Fuckel and fitting the data of the protologue. The collection
Type. Lectotype (designated here, MBT198518): Germany, c. Budenheim, UPS F-127433 contains three basidiomata, all of them well-
Fuckel 317, UPS F-127434! conserved and two of them with the pseudoparenchymatous
layer in good condition, glued to a sheet of paper in different
= G. queletii Hazsl., Verh. Bot. Vereins Prov. Brandenburg 24: 136. 1883,
positions, and clearly showing both big crystals on endoperidial
‘Geaster Quéletii ’. — Type: Lectotype (designated here, MBT198519):
France, Jura, Quélet, S F16394!
surfaces, and a whitish stalk. The collection UPS F-127434
consists of a single, well-conserved basidioma with big crys-
Synoptic description (based on Staněk (1958, as G. minimum tals on the endoperidial surface, being the stalk hidden by the
var. minimum) and studied specimens) — Exoperidium (9–)11– pressing procedure, but if the endoperidial body is carefully
42(–60) mm diam in horizontal position, arched, with 5–13(–14) lifted, a short, whitish stalk can be seen. This collection was
not hygrometric rays. Endoperidial body 3 –12 (–15.5) mm additionally marked by C.G. Lloyd as “This is a type but the spe-
diam, cream coloured, greyish or brownish, less frequently cies = G. minimus”, and the number 317 is the one that Fuckel
whitish. Mesoperidium as a dense layer of crystalline matter. (1860) used for enumerating the species in the protologue.
Largest mesoperidial crystals of COD on the endoperidial sur- Finally, the collection Herbier Barbey-Boissier 2138 contains
face 65 –165 µm diam, bipyramidal, normally very abundant. another three basidiomata, one of them rather young and the
Largest mesoperidial crystalline aggregates of COM scales other two somewhat old, showing big crystals on endoperidial
(7–)10 – 62(–120) µm diam, often present but inconspicuous, surfaces and whitish stalks as well. All these specimens are in
sometimes abundant. Peristome fibrillose, flat or conical, mostly full agreement with the concept of the species followed here.
distinctly delimited. Stalk more or less stout, (0.3–)0.5–2.0 mm One of these collections can be selected as lectotype and, since
high, whitish to cream coloured. Basidiospores globose, 4.5– Lloyd considered the collection UPS F-127434 as ‘type’, and
6.5(–7.5) µm diam, with (0.3 –)0.4 – 0.8(–1.0) µm high warts, the number 317 is the one used by Fuckel in the protologue of
ornamentation verrucose to irregularly pilate. Broadest capil the species, we designate this collection as the lectotype for
litial hyphae 4.5 – 9.5 µm wide. Pseudoparenchymatous layer the species. Geastrum queletii was briefly described by Hazs-
of thin-walled (mostly ≤ 1 µm thick) cells. linszky (1883) based on material collected by Quélet (1873),
Ecology & Distribution — Confirmed collections of this who named it as ‘Geaster umbilicatus’. There is one specimen
species came from calcareous soils on both open places and at the Stockholm herbarium marked as ‘typus’ (S F16394) by
forests, of a wide range of biomes (‘Boreal forests /taiga’, a latter hand than that of Hazslinsky, and in the original label
‘Temperate conifer forests’, ‘Temperate broadleaf and mixed it is marked ‘Geaster Quéleti Hasz.’. It consists of a single,
well-preserved basidioma, which shows big crystals on the lumen normally wide, often sinuous, narrowing at base and
endoperidial surface and a whitish stalk, collected from Jura by apex, tips often acute. Mesoperidium present on the endoper-
Quélet. This specimen has been designated as lectotype and, in idium and pseudoparenchymatous layer surfaces, consisting of
agreement with our concept of G. granulosum, it is considered rather large bipyramidal crystals of COD, the largest 70–135 µm
as a heterotypic synonym. diam, and abundant 1.5–4.0 µm wide, hyaline to pale brownish,
branched, thin-walled, clamped hyphae; rounded crystalline
Geastrum kuharii J.C. Zamora, sp. nov. — MycoBank aggregates of COM scales up to 70–95 µm diam, sometimes
MB810503; Fig. 4, 10e inconspicuous. Pseudoparenchymatous layer of thin-walled
(mostly ≤ 1 µm thick), hyaline cells, variable in shape and size,
Type. Argentina, Buenos Aires, Partido de Berazategui, Pereyra, Pereyra
about 15–80 × 10–42 µm diam. Fibrous layer with 1.5–6.0
Iraola park, Pinus sp., 16 June 2012, F. Kuhar, V. Castiglia & J.C. Zamora,
holotype MA-Fungi 83795!, isotypes in AH 45202! and UPS!
µm wide, hyaline to very pale yellowish, aseptate, straight or
slightly sinuous, intertwined, mostly unbranched, comparatively
Etymology. The specific epithet is dedicated to Francisco Kuhar, Argentin- thick-walled (0.7–2.5 µm thick) hyphae, lumen visible. Mycelial
ian mycologist expert in Geastrum, who helped collecting the type specimen.
layer double-layered; inner layer consisting of 1.5 – 3.0 µm
Diagnosis — Exoperidium 45 – 62 mm diam in horizontal posi- wide, strongly glued together, more or less hyaline, branched,
tion, arched, with 8 –11 not hygrometric rays. Endoperidial thin-walled and clamped hyphae; outer layer with 1.0–4.0 µm
body 13 – 21 mm diam, greyish cream to brownish grey. Meso- wide, hyaline to somewhat yellowish, aseptate, rarely branched,
peridium rather well-developed, as a mealy cover with big comparatively thick-walled (0.5–2.0 µm thick) hyphae, lumen
crystals. Largest mesoperidial crystals of COD on the endo very narrow and difficult to perceive. Rhizomorphs with bipy-
peridial surface 75 –130 µm diam, bipyramidal. Largest ramidal crystals of COD, often grouped in rose-like aggregates.
mesoperidial crystalline aggregates of COM scales 70 –95 µm Ecology & Distribution — Only known from humic soils of
diam, sometimes not very conspicuous. Peristome fibrillose, the ‘Temperate grasslands, savannas and shrublands’ biome
flat, well-delimited. Stalk stout 1.5 – 2.5 mm high, pale brown of the Neotropic ecozone.
to brown. Basidiospores globose, 4.5 – 5.5(– 6.0) µm diam, Additional specimens examined (paratypes). Argentina, Buenos Aires,
with 0.4 – 0.7(– 0.8) µm high warts. Broadest capillitial hyphae Partido de Berazategui, Pereyra, Pereyra Iraola park, Pinus sp., 16 June
6.0–7.0 µm wide, ornamentation verrucose to irregularly pilate. 2012, F. Kuhar, V. Castiglia & J.C. Zamora, MA-Fungi 86914; Entre Ríos,
Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm Colón, Ubajay, marginal forest, 2 June 2012, S. Suaza & J.C. Zamora, MA-
thick) cells. Rhizomorphs with bipyramidal crystals of COD, Fungi 86913.
grouped in rose-like aggregates.
Macroscopic characteristics — Unexpanded basidiomata Geastrum marginatum Vittad., Monographia Lycoperdineo-
20–33 mm diam, subglobose to depressed, with a rounded rum: 20. 1842, ‘Geaster marginatus’ — Fig. 6, 11e
apex or a flat umbo. Exoperidium splitting in 8 –11 more or less Type. Lectotype (designated here, MBT198520): f. 6 of pl. I in Vittadini
equal to unequal rays, 30 – 55 mm diam apparently, 45 – 62 (1842). Epitype (designated here, MBT198521) that supports the lectotype
mm diam when forced in horizontal position, arched, not hy- cited above: Spain, Madrid, El Pardo-Fuencarral, encinar, 6 Dec. 2000, J.C.
grometric. Mycelial layer rather thick, whitish to pale cream, Campos y familia, MA-Fungi 48129!
sometimes with pinkish shades when fresh, strongly intermixed = Geastrum cesatii Rabenh., Bot. Zeitung (Berlin) 9, 36: 628. 1851,
with debris from the substrate, more or less strongly adhered ‘Geaster Cesatii ’. — Type: Unknown (but see nomenclatural notes below).
to the fibrous layer but often peeling-off in some parts. Fibrous = Geastrum minimum var. fumosicollum V.J. Staněk in Pilát, Flora ČSR
layer more or less coriaceous when denuded, pale-cream to B-1: 435, 786. 1958. — Type: Czech Republic, Bohemia, Velká Chuchle ap.
cream, in fresh sometimes turning pinkish when damaged. Praha, 18-IX-1954, E. Wichanský, holotype PRM 842884! Various paratypes
were also cited by Staněk (1958).
Pseudoparenchymatous layer whitish at first, surface cream to
?= Geastrum juniperinum T. Macbr., Mycologia 4, 2: 85. 1912, ‘Geaster
greyish, normally superficially cracked, < 0.5 mm thick in dry juniperinus’. — Type: Unknown. In absence of original specimens, f. 1 of pl.
state, about 2.0 – 4.0 mm thick when fresh, not very persistent. LXII in Macbride (1912) can be selected as lectotype.
Endoperidial body globose to subglobose, 13 – 21 mm diam,
endoperidial surface glabrous, brownish. Mesoperidium con- Synoptic description (based on Staněk (1958, as G. minimum
spicuous, with both abundant generative hyphae and crystal- var. fumosicollum) and studied specimens) — Exoperidium
line matter. Peristome fibrillose, often slightly darker than the 13–40 mm diam in horizontal position, arched, with 5–13(–14)
endoperidial surface, flat, thickened, and distinctly delimited. not hygrometric rays. Endoperidial body 3–14 mm diam, greyish
Stalk stout, 1.5 – 2.5 mm high, pale brown to brown. Apophysis to greyish brown, rarely whitish or very dark. Mesoperidium as
present, well-developed or not, sometimes almost ring-like, with a thin layer of sparse pruina. Largest mesoperidial crystals of
the same colour as the endoperidium. Columella intruding about COD on the endoperidial surface (5–)7.5–70(–95) µm diam,
1/2 into the glebal mass, stout, with a rounded apex. Mature bipyramidal, often rather scarce. Largest mesoperidial crystal
gleba dark brown. line aggregates of COM scales 5–25(–40) µm diam, normally
Microscopic characteristics — Basidia narrowly ellipsoid to inconspicuous, rarely abundant. Peristome fibrillose, flat or
subcylindric or more or less lageniform, 15 – 26 × 4.5 – 6.5 µm, conical, mostly distinctly delimited. Stalk more or less stout,
with (4 –)5 – 8 short sterigmata. Basidiospores globose to sub- (0.3–)0.5–2.0 mm high, mostly brownish, to greyish brown or
globose, 4.5–5.5(–6.0) µm diam, brownish, with 0.4–0.7(–0.8) even blackish, less frequently pale coloured. Basidiospores
µm high brown warts, verrucose to irregularly pilate ornamenta- globose, 4.5–7.0(–8.0) µm diam, with (0.3–)0.4–0.8(–1.0) µm
tion. Broadest capillitial hyphae 6 –7 µm wide, aseptate, very high warts, ornamentation verrucose to irregularly pilate. Broad
rarely branched, normally straight, thick-walled (1.5 – 3 µm est capillitial hyphae 4.5–9.0 µm wide. Pseudoparenchymatous
thick), with narrow lumen, mostly visible; tips acute to rounded; layer of thin-walled (mostly ≤ 1 µm thick) cells.
surface often covered with debris. Endoperidium composed of Ecology & Distribution — Confirmed specimens came from
1.5–6.0 µm wide, pale yellowish to yellowish brown, aseptate, mostly sandy, siliceous or slightly calcareous soils, from both
mostly unbranched, straight to slightly sinuous, strongly inter- open places and forests, of several biomes (‘Temperate conifer
twined, thick-walled hyphae, lumen visible; protruding hyphae forests’, ‘Temperate broadleaf and mixed forests’, and ‘Medi-
not seen. Peristomal hyphae 5.0 –10.0 µm wide, brownish, terranean forests, woodlands and scrub’) of the Neartic and
aseptate, mostly unbranched, thick-walled (1.0 – 3.0 µm thick), Paleartic ecozones.
Additional specimens examined. Italy, Ne’ luoghi arenosi e scarsi di erbe Geastrum quadrifidum Pers., Neues Mag. Bot. 1: 86. 1794;
lungo gli argini del fume Sesia presso Vercelli, Dec. 1859–60, Cesati, Erbario Pers., Syn. Meth. Fung.: 133. 1801. — Fig. 3, 11f
Crittogamico Italiano ser. I, B 70 0014089; ibid., ex herb. Sydow, S; Sesia
presso Vercelli, 1860, Cesati, S. – Spain, Ávila, camino de Piedralaves a La Type. Lectotype (designated here, MBT198522): f. 1 of pl. XXXVII in
Adrada, 4 Dec. 1984, F.D. Calonge, MA-Fungi 8112; Huesca, Javierregai, Schmidel (1793).
676 m, encinar con jaras, suelo arcilloso, 28 Oct. 2005, F. Prieto, Á. González
& C. Diego, herb. Zamora 108; Canary Islands, La Gomera, pantano de = Geastrum coronatum (Scop.) J. Schröt. in Cohn, Krypt.-Fl. Schlesien
Arure-Acardece, 882 m, en zona arbustiva, 26 Nov. 2012, J. Fernández, 3 –1, 6: 702. 1889, ‘Geaster coronatus’, nom. illeg. (Art. 53.1).
M. Oyarzabal, F. Hidalgo & R. Martínez, ERRO 2012112609; Jaén, Baños ≡ Lycoperdon coronatum Scop., Fl. Carniol. 2: 490. 1772. — Type:
de la Encina, junto al río Aliseda, 28 Nov. 1993, F.D. Calonge, MA-Fungi Lectotype (designated here, MBT198523): f. 3 of pl. CLXXXIII in Schaeffer
32395; Jaén, Venta de los Santos, 12 Oct. 2000, J.C. Zamora, MA-Fungi (1763).
48128; Madrid, Majadahonda, 3 Dec. 1989, F.D. Calonge, MA-Fungi 31530; = Geastrum quadrifidum var. minus Pers., Syn. Meth. Fung.: 133. 1801.
Madrid, El Pardo, entrada por El Goloso, en suelo arenoso de encinar, 11 ≡ Geastrum minus (Pers.) G. Cunn., Proc. Linn. Soc. New South Wales
Nov. 1978, E. Álvarez, MA-Fungi 724; Madrid, El Pardo, bajo encina, 4 Feb. 51: 81. 1926, ‘Geaster ’. — Type: Lectotype (designated here, MBT198524):
1979, E. Álvarez, MA-Fungi 725; Salamanca, Valdelosa, pradera al borde f. 2 of pl. XXXVII in Schmidel (1793).
de camino, 10 Oct. 1993, J. Lozano, MA-Fungi 32394; Sevilla, Puebla de = Geastrum quadrifidum var. sabulosum V.J. Staněk in Pilát, Flora ČSR
los Infantes, Los Cerrillares, 8 Dec. 2008, M.Á. Ribes, herb. Ribes 081208- B-1: 439, 794. 1958. — Type: Slovakia, Píscové přesypy mladém boru u
03; Teruel, Albarracín, en encinar, 26 Oct. 1988, F.D. Calonge, MA-Fungi Malacek, 1 May 1954, V.J. Staňkovi, holotype PRM (photo!); Kúty, loco
21695. – Sweden, Gotland, Holmhällar (close to the hotel), 28 Sept. 2011, arenoso, 17 Nov. 1955, F. Kotlaba, paratype PRM 842888!
open area on sandy soil, among lichens, mosses and grasses, not far
away from Pinus sylvestris, J.C. Zamora, M. Jeppson & M. Lathi, MA-Fungi Synoptic description (based on Sunhede (1989) and studied
86669. – USA, Arizona, Coconino Co., Walnut Canyon National Monument, specimens) — Exoperidium (15–)17–50(–90) mm diam in hori-
N35°10" W111°30.26", Juniperus, 12 Oct. 1986, J. States AEF 528, MICH zontal position, arched, with (3–) 4–5(–6) not hygrometric rays.
28120; Arizona, Coconico Co., Walnut Canyon National Monument, in open Endoperidial body (3.5 –)5 –12(–16) mm diam, pale cream
area around debris, Pinus edulis and Juniperus, 17 Oct. 1986, J. States AEF to rather dark greyish brown. Mesoperidium as a thin layer of
518, MICH 28119b.
pruina. Largest mesoperidial crystals of COD on the endoperi
Nomenclatural notes — There is one collection at herbarium dial surface (10–)15–50(–60) µm diam, bipyramidal, abundant.
K marked by a latter hand than that of Vittadini (1842) as Mesoperidial crystalline aggregates of COM scales normally
“Geaster marginatum Vittad. Isotypus. Overseas, sine loc.; on very scarce or inconspicuous, about 5–25(–30) µm diam. Peri
soil; ex herb. M. J. Berkeley; Herbarium Mycologicum Berke- stome fibrillose, conical to papillate, rarely flat, mostly sharply
leyanum, presented by the Rev. M.J. Berkeley, 1879; K(M) delimited. Stalk often stout, (0.3 –)1.0 – 2.0(– 2.5) mm high,
169190”, which consists of three more or less well-preserved whitish to brownish grey. Basidiospores globose, (4.5–)5.0–
basidiomata, in agreement with the protologue and with the 6.0(–6.5) µm diam, with (0.3–)0.4–0.8 µm high warts, orna-
concept of the species presented here. However, whether it is mentation verrucose to irregularly pilate. Broadest capillitial
part of the original material studied by Vittadini or not is difficult hyphae (4.0–)4.5–9.5 µm wide. Pseudoparenchymatous layer
to say with certainty, due to the absence of ecological and of thin-walled (mostly ≤ 1 µm thick) cells.
geographical data on the herbarium label. The description and Ecology & Distribution — Mainly growing in calcareous
the iconography from Vittadini (1842) suggest that the concept soils of ‘Boreal forests/taiga’, ‘Temperate conifer forests’, and
he used may be the same as considered here, but they are ‘Temperate broadleaf and mixed forests’ biomes of the Neartic
not detailed enough to clearly distinguish this species from the and Paleartic ecozones, but also present in the ‘Mediterranean
morphologically close G. granulosum, because crystals on the forests, woodlands and scrub’ biome (Sunhede 1989, Calonge
endoperidial surface and the stalk colour are not mentioned. 1998, Bates 2004).
For this reason, although f. VI of t. I in Vittadini (1842) is part of Additional specimens examined. Spain, Cuenca, Almodóvar del Pinar,
the protologue and has been designated here as lectotype, in bosque de Q. rotundifolia, Q. coccifera, Pinus pinaster y algún Q. faginea,
absence of unambiguous original material, an epitype should suelo calizo, 25 Oct. 2008, J.F. Mateo, F. Gracia & J.C. Campos, herb. Zamora
also be chosen in order to fix the interpretation of this species. 300; Huesca, Canfranc, bajo Picea abies, 14 Oct. 2006, M. Chiaffi, herb.
Therefore, we have designated as epitype one sequenced Zamora 170; Huesca, Bielsa, valle de Pineta, bajo Abies alba, 13 Oct. 2012,
J. Hernanz, herb. Zamora 561; Madrid, Guadalix de la Sierra, ‘El Montecillo’,
specimen from the Mediterranean basin, collected in sandy,
bosque de Quercus rotundifolia y Q. faginea, suelo calizo, 5 Nov. 2011, A. Díaz,
siliceous soil, which consists of six well-developed and typical herb. Zamora 590; Orense, O Barco de Valdeorras, pinar Vegamolinos,
basidiomata. Geastrum cesatii (Rabenhorst 1851) was de- bajo Pinus sp., entre musgos, 11 Jan. 2006, C. Ruiz, herb. Zamora 139. –
scribed from specimens collected by Cesati in Piedmont (near Sweden, Gotland, Stenkyrka parish, Snipklint, Picea abies, Pinus sylvestris
Vercelli). We have not been able to trace any original material, forest, with Juniperus communis on limestone, on needle litter under dense
but there are two specimens in the Stockholm herbarium (S), spruces, 7 Apr. 1968, D. Lowgren, B. Lowgren & S. Sunhede, herb. Sunhede
and one in the Berlin herbarium (B), collected by Cesati near 97a; Uppland, Älkarleby, Billuddens naturreservat, under Pinus sylvestris,
Picea abies, and Betula pendula, on sandy, calcareous soil, 15 Oct. 2011,
Vercelli after Rabenhorst’s (1851) publication, being part of
I. Olariaga & J.C. Zamora, MA-Fungi 86671. – USA, California, Siskiyou
the Erbario Crittogamico Italiano serie I, n. 590, in agreement Co., 2 mi E of McCloud, in duff in Pseudotsuga menziesii, Abies concolor,
with the protologue and with the concept of G. marginatum Quercus and Pinus forest, 25 Aug. 1990, Richter D. DR90-05, MICH 39032;
followed here. Thus, the name is considered as a heterotypic Colorado, Boulder Co., vicinity Castle Rock in Middle Boulder Creek Canyon,
synonym of G. marginatum. In absence of original material, 1.5 mi northeast of Tungsten, N39°59' W105°27', c. 7900 ft., 11 Sept. 1970,
any of these collections may serve as a good neotype for the R. Fogel F2078, MICH 72512; Minnesota, Clearwater Co., Itasca St. Pk.,
name. Geastrum minimum var. fumosicollum (Staněk 1958) is 26 Sept. 1970, K.E. Harrison, MICH 72511.
also considered as a heterotypic synonym, since morphological Nomenclatural notes — As noted by Sunhede (1989), the
data agree with those of typical basidiomata of G. marginatum, specimen designated as type by Eyndhoven (1937) should
and molecular data place the holotype within the strongly sup- be considered as a neotype. Therefore, although in perfect
ported G. marginatum clade (Fig. 1). Geastrum juniperinum agreement with the current concept of this species, it is to be
(Macbride 1912) was not described in detail, but according to superseded according to Art. 9.19(a) because original material
the iconography of the protologue it could be possibly another (pl. XXXVII in Schmidel (1793)) exists. Protologues and original
heterotypic synonym. Since original specimens have not been material of G. coronatum (Scop.) J. Schröt. (nom. illeg.), G. qua-
studied by us yet, and considering that some species in G. sub- drifidum var. minus, and G. quadrifidum var. sabulosum agree
sect. Quadrifida are difficult to be distinguished, we prefer to with the concept of the species followed here, and are consid-
indicate such synonymy with an expression of doubt. ered heterotypic synonyms.
Geastrum subsect. Sulcostomata V.J. Staněk in Pilát, Flora 9. Columella mostly with a persistent and pointed apex; South
ČSR B1, Gasteromycetes: 781. 1958, nom. inval. (Art. 22.2) American species . . . . . . . . . . . . . . . . . . . G. glaucescens
9. Columella mostly with a deciduous, often rounded but some-
≡ Geastrum subg. Pectinata Dörfelt & Müll.-Uri in Dörfelt, Die Erdsterne:
Geastraceae und Astraeaceae: 17. 1985. — Type: Geastrum pectinatum times pointed, apex; South European species . . . . . . . . . .
Pers. (Fig. 1E). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. parvistriatum
= Geastrum sect. Striata De Toni, Rev. Mycol. 9, 34: 64. 1887, ‘Striati ’.
— Type: Geastrum striatum DC.
Geastrum biplicatum Berk. & M.A. Curtis, Proc. Amer. Acad.
Nomenclatural notes — This name is currently not validly Arts 4: 124. 1860, ‘Geaster biplicatus’ — Fig. 12a
published according to Art. 22.2 (McNeill et al. 2012) but, as Type. Lectotype (designated here, MBT198525): Japan, Bonin Islands, on
noted in Zamora et al. (2014) by the nomenclature editor of the the ground, US N Pacific Expedition Exp. (ex. herb. Hooker), K(M) 180394!
journal Taxon, the name is used pending potential amendments (probably one of various syntypes).
to Art. 22.2.
Synoptic description (based on the type specimen only) —
Exoperidium 34 mm diam in horizontal position, with 7 non-
Key to species in Geastrum subsect. Sulcostomata
hygrometric rays. Endoperidial body 11 mm diam, dark brown.
1. Apophysis very well-developed, solid, ring-like . . . . . . . . . 2 Mesoperidium well-formed, powdery. Largest mesoperidial
1. Apophysis different or absent; if present, then as a fold of crystals of COD on the endoperidial surface about 15 µm diam,
the endoperidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 bipyramidal. Peristome sulcate, more or less distinctly delimited,
with about 19 folds. Stalk slender, about 2.0 mm high and 1.0
2. Apophysis with a sharp and acute edge, basidiospores 5.0– mm wide, brownish. Base of the endoperidial body more or
6.0 µm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . G. striatum less sulcate. Basidiospores globose, (4.5–)5.0–6.0 µm diam,
2. Apophysis with a rounded edge, basidiospores 4.5 –5.0 µm with 0.5 –1.0 µm high warts, ornamentation baculate-pilate.
diam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. aff. striatum Broadest capillitial hyphae 10.5 µm diam. Cells of the pseudo
3. Stalk (2.5–)3–13 mm high, frequently slender, length/breath parenchymatous layer deteriorated and thus not studied, ex
= 0.7–7.5(–8.6); species preferring humid to subhumid habi- pectedly thin-walled.
tats; basidiospore ornamentation clearly baculate-pilate . 4 Ecology & Distribution — Insufficiently known; the type
3. Stalk 0.5–2.5(–3) mm high, mostly stout, length /breath = 0.5– specimen was collected in the Bonin Islands (Berkeley & Curtis
1(–1.5); species preferring subhumid to semixeric habitats; 1860), which are located in the ‘Tropical and subtropical moist
basidiospore ornamentation verrucose to irregularly pilate broadleaf forests’ biome of the Oceania ecozone.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Additional remarks — Specimens included in the phyloge-
4. Largest mesoperidial crystals 20–55 µm diam; Australasian netic tree as ‘G. cf. biplicatum’ were not available for morpho-
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. tenuipes logical study even if requested on loan (see Discussion).
4. Mesoperidial crystals up to 15(– 32) µm diam; not Australa-
sian species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Geastrum glaucescens Speg., Anales Mus. Nac. Hist. Nat.
5. Basidiospores (4.5 –)5.0 – 6.5 µm diam; stalk 0.7– 2.5 mm
Buenos Aires 23: 14. 1912, ‘Geaster ’ — Fig. 7, 13a
width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5. Basidiospores 5.5 –7.5(– 8.5) µm diam; stalk (1.6 –)2.0 – Type. Argentina, Buenos Aires, La Plata, Spegazzini, Sept. 1894, holotype
8.0(– 9.0) mm width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 LPS 15860!
6. Broadest capillitial hyphae 6.0 – 8.0 µm diam; Pantropical Synoptic description — Exoperidium 18–45 mm diam in hori-
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. plicatum zontal position, arched, divided in 6–11 non-hygrometric rays.
6. Broadest capillitial hyphae up to 10.5 µm diam; Oceanian Endoperidial body 5–16 mm diam, mostly dark brown to black-
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. biplicatum ish. Columella with a persistent and pointed apex, rarely
7. Broadest capillitial hyphae (5.5 –)6.0 – 9.0(– 9.5) µm diam; rounded. Mesoperidium very abundant, powdery. Largest meso
stalk relatively stout, length /breath = 0.6 – 2.3, light or dark peridial crystals of COD on the endoperidial surface 6.0–15 µm
coloured; peristome with (17–)22 – 52(– 60) folds; endoperi diam, bipyramidal. Peristome sulcate, distinctly delimited or not,
dium mostly light coloured, from cream to brownish grey, with 11–25 folds. Stalk 0.5–2.5 mm high, mostly brownish to
never blackish; Mediterranean and Macaronesian species dark brown. Basidiospores globose, 5.0–7.0(–7.5) µm diam,
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . G. meridionale with 0.5–1.0(–1.2) µm high warts, ornamentation verrucose to
7. Broadest capillitial hyphae (8.0–)9.0–13.0(–15.0) µm diam; irregularly pilate. Broadest capillitial hyphae 6.0–9.0 µm diam.
stalk mostly slender, length/breath = 1.7– 4.0, mostly light Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm
coloured; peristome with 11– 35 folds; endoperidium very thick) cells.
variable in colour, often dark, even blackish; Temperate to Ecology & Distribution — This species grows mostly under
Boreal species . . . . . . . . . . . . . . . . . . . . . . . G. pectinatum mesquite (Prosopis spp.), and has been found in ‘Temperate
grasslands, savannas and shrublands’ and ‘Tropical and sub-
8. Basidiospores 4.0 – 5.5 µm diam, brownish, ornamentation
tropical grasslands, savannas and shrublands’ biomes of the
0.3 – 0.6 µm high; broadest capillitial hyphae 4.5 –6.0 µm
Neotropic ecozone. Currently only known from Argentina (Soto
diam; glebal mass brownish to dark brown; basidiomata rather
& Wright 2000, Kuhar et al. 2013).
slender, not reminiscent G. striatum, endoperidial body 3.5–
6.5 mm diam, exoperidium 12 – 24 mm diam . G. papinuttii Additional specimens examined. Argentina, Catamarca, San Fernando
8. Basidiospores (4.5–)5.0–7.0 µm diam, dark brown to blackish, del Valle de Catamarca, Prosopis sp., 10 Apr. 2012, L. Papinutti & J.C.
Zamora, MA-Fungi 83763; La Rioja, Castro Barros, Anjullón, Prosopis sp.,
ornamentation (0.4–)0.5–1.0(–1.2) µm high; broadest capil-
8 Apr. 2012, L. Papinutti & J.C. Zamora, MA-Fungi 83762; ibid., MA-Fungi
litial hyphae 5.5–9.0 µm diam; gleba dark brown to blackish; 86906; ibid., MA-Fungi 86907; ibid., MA-Fungi 86908; ibid., MA-Fungi 86909;
basidiomata comparatively stout, often strongly reminiscent ibid., MA-Fungi 86910; La Rioja, Santa Cruz, S28°28'35.11" W67°41'54.45",
G. striatum, endoperidial body 5 –16 mm diam, exoperidium on abundant Prosopis sp. leaf litter, 27 Mar. 2008, L. Papinutti & G. Rolón,
(13 –)18 – 47 mm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 BAFC 51940.
Geastrum meridionale J.C. Zamora, sp. nov. — MycoBank Etymology. The epithet refers to the southern distribution of the species
MB810504; Fig. 12b in Europe, especially when compared with the mostly central and northern
distribution of its closest relative, G. pectinatum.
Type. Spain, Madrid, Ciudad Universitaria, E.T.S.I. de Montes, under
Calocedrus decurrens, 17 Oct. 2007, J.C. Zamora 252, holotype MA-Fungi Diagnosis — Exoperidium 45–125 mm diam in horizontal posi-
87325!, isotypes in AH 45203! and UPS! tion, divided in 5–12 non-hygrometric rays. Endoperidial body
12–33 mm diam, cream to brownish grey, rarely dark brown.
Mesoperidium very abundant, powdery. Largest mesoperidial
Fig. 12   Morphological characters in Geastrum subsect. Sulcostomata pro parte (G. pectinatum group). a. G. biplicatum (lectotype); b. G. meridionale (holo-
type); c. G. pectinatum (epitype); d. G. cf. plicatum (MA-Fungi 83774); e. G. tenuipes (CANB 738350). — a1, b1, c1, d1, e1. Basidiomata habit; a2, b2, c2, d2,
e2. detail of the peristome; a3, b3, c3, d3, e3. detail of the stalk and apophysis; a4, b4, c4, d4, e4. mesoperidial crystals of COD on the endoperidial surface;
a5, b5, c5, d5, e5. basidiospores. — Scale bars: a1, b1, c1, d1, e1 = 5 mm; a2, b2, c2, d2, e2, a3, b3, c3, d3, e3 = 2 mm; a4, b4, c4, d4, e4 = 10 µm; a5, b5,
c5, d5, e5 = 2 µm.
crystals of COD on the endoperidial surface 6 –13 µm diam, Ecology & Distribution — This species grows on both calcar-
bipyramidal. Peristome sulcate, distinctly delimited or not, with eous and siliceous soils, not rarely in anthropized environments,
about (17–) 22 – 52(– 60) folds. Stalk stout to more or less slen- and seems to be characteristic of the ‘Mediterranean forests,
der, (2.5–)3.0–9.0(–9.5) mm high and 2.0–8.0(–9.0) mm wide, woodlands and scrub’ biome of the Paleartic ecozone.
cream to rather dark brownish grey. Base of the endoperidial
Additional specimens examined (paratypes). Portugal, Estremadura,
body from smooth to plicate or conspicuously sulcate. Basi Mata do Solitario, 12 Nov. 1991, F.D. Calonge, MA-Fungi 31164. – Spain,
diospores globose, (5.5 –)6.0 –7.5(– 8.5) µm diam, with 0.6– Cáceres, Almaraz, mediterranean woodland, calcareous soil, 17 Jan. 2014,
1.5(–1.6) µm high warts, ornamentation baculate-pilate. Broad C. Gelpi, herb. Zamora 617; Cádiz, Grazalema, Llanos del Campo, 30STF8170,
est capillitial hyphae (5.5 –) 6.0 – 9.0(– 9.5) µm diam. Pseudo- 760 m, bosque mixto Quercus faginea y Ceratonia siliqua, 15 Dec. 2003,
parenchymatous layer of thin-walled (mostly ≤ 1 µm thick) cells. A. Castro 688, MA-Fungi 59644; Canary Islands, Tenerife, Las Mercedes,
Cruz del Carmen, 926 m, en bosque de layal-brezal, 30 Dec. 2007, D. Cha
Macroscopic characteristics — Unexpanded basidiomata vez, V. Escobio, J. Caridad & M.Á. Ribes, herb. Ribes 301207-43; Córdoba,
up to 30 mm diam, subglobose, with a rounded apex or a flat Cabra, encinar, 4 Nov. 1990, unknown legit, AH 12674; Córdoba, arroyo de
umbo. Exoperidium splitting in 5 –12 more or less equal to Pedroche, UG4499, bajo Quercus rotundifolia, 24 Dec. 1999, A. Castro 6,
MA-Fungi 55189; Guadalajara, embalse de El Vado, under Pinus nigra,
unequal rays, 30 –107 mm diam apparently, 45 –125 mm diam
Cupressus sp., Quercus pyrenaica, Q. rotundifolia, and Gleditsia triacanthos,
when forced in horizontal position, arched, often more or less calcareous soil, 6 Dec. 2007, M. Bueno & A. Zapata, herb. Zamora 276; ibid.,
hygrometric. Mycelial layer rather thick, whitish to pale cream under Pinus sp., Cistus sp., Quercus sp. and Cupressaceae, 12 Dec. 2012,
or somewhat yellowish, strongly intermixed with debris from M. Bueno & A. Zapata, herb. Zamora 465; Madrid, Ciudad Universitaria,
the substrate, more or less strongly adhered to the fibrous E.T.S.I. de Montes, under Cedrus libani, 13 Nov. 2004, L. Rubio, herb. Zamora
layer but sometimes peeling-off in some parts. Fibrous layer 65; ibid., under Cedrus sp., 20 Aug. 2005, J.C. Campos & J. Daniel-Arranz,
herb. Zamora 82; ibid., under Cupressus sp., 22 Nov. 2005, J.C. Zamora, MA-
papyraceous to coriaceous when denuded, whitish to cream
Fungi 63558 (duplo herb. Zamora 118); ibid., under Cupressus sp., 10 July
coloured. Pseudoparenchymatous layer whitish to greyish 2006, J.C. Zamora, herb. Zamora 153; ibid., under Cedrus libani, 30 Oct.
cream, not or only superficially cracked, < 0.5 mm thick in dry 2006, J.C. Zamora, herb. Zamora 185; ibid., under Thuja sp., 23 Nov. 2006,
state, about 1 mm thick when fresh, rather persistent. Endope J.C. Zamora, herb. Zamora 192; ibid., under Calocedrus decurrens, 17 Oct.
ridial body globose, subglobose, discoid or broadly ellipsoid, 2007, J.C. Zamora, herb. Zamora 262; ibid., under Cedrus sp., 15 Oct. 2008,
rarely irregular, 12–33 mm diam, cream to brownish grey, rarely J.C. Zamora, herb. Zamora 294; ibid., under Eucalyptus sp. and Cupressus
dark brown; endoperidial surface glabrous. Mesoperidium as a sempervirens, 15 Oct. 2008, J.C. Zamora, herb. Zamora 295; ibid., under
Cedrus deodara, 14 Nov. 2008, L. Rubio Casas, AH 42179; ibid., under
very thick, powdery to creamy, layer of whitish, pale brownish
Chamaecyparis lawsoniana, 20 Nov. 2008, J.C. Zamora, herb. Zamora 315;
or greyish pruina. Peristome sulcate, with the same colour as ibid., under Cupressus sempervirens and Mahonia aquifolium, 11 Nov. 2011,
the endoperidial surface, narrowly to broadly conical, distinctly J.C. Zamora, herb. Zamora 587; ibid., under Eucalyptus sp. and Cupressus
or indistinctly delimited. Stalk present, stout to more or less sempervirens, 11 Nov. 2011, J.C. Zamora, herb. Zamora 588; Madrid, parque
slender, (2.5 –)3.0 – 9.0(– 9.5) mm high, cream to rather dark Pedriza de Manzanares, pinar, 14 Oct. 1995, F.D. Calonge, MA-Fungi 33803;
brownish grey. Apophysis present or absent, with the same Madrid, San Agustín de Guadalix, suelo húmico de encinar, 4 Dec. 2004,
colour as the endoperidium or slightly darker, smooth to plicate J.C. Campos, MA-Fungi 60901; Madrid, Villaviciosa de Odón, cerca de la
Residencia de Ancianos de la C.A.M., bajo Quercus rotundifolia, Oct. 1998,
or conspicuously sulcate. Columella intruding beyond the mid-
J.C. Zamora, herb. Zamora 1; Mallorca, Artá, 15 Oct. 1990, C. Constantino,
dle into the glebal mass, more or less fusiform, with an acute MA-Fungi 27661; Mallorca, Esporles, Port d’es Canonge, 22 Nov. 1987,
apex. Mature gleba brownish to dark greyish brown. L. Siquier 1 V, MA-Fungi 20651; Mallorca, Predio ca na Magdalena Noia, Pol-
Microscopic characteristics — Basidia not seen. Basidio lensa, J.L. Siquier, MA-Fungi 20834; Mallorca, Son Forté, Artá, 9 Nov. 1991,
spores globose to subglobose, (5.5 –)6.0 –7.5(– 8.5) µm diam, J.L. Siquier, MA-Fungi 27662; Palencia, Ampudia, bosque de Cupressus
brownish to yellowish brown, with 0.6–1.5(–1.6) µm high brown arizonica, 26 Jan. 2008, A. García & M. Sanz, AVM 2856; ibid., AVM 2857;
Toledo, Quercus rotundifolia, 4 Dec. 2007, J. De Esteban, herb. Zamora
warts, ornamentation baculate-pilate. Broadest capillitial hyphae
616; Valladolid, parque de las Contiendas, bajo Cupressus arizonica, 28
(5.5–)6.0–9.0(–9.5) µm diam, aseptate, very rarely branched, nor- Mar. 2011, A. García AVM 2708.
mally straight, thick-walled (2.0 – 4.0 µm thick), with narrow
lumen but mostly visible; tips acute to rounded; surface often
covered or not with debris. Endoperidium composed of 2.0–5.5 Geastrum papinuttii J.C. Zamora, sp. nov. — MycoBank
µm wide, pale yellowish to yellowish brown, aseptate, mostly MB810505; Fig. 7, 13c
unbranched, slightly sinuous, strongly intertwined, thick-walled Type. Argentina, Santiago del Estero, Guasayán, Prosopis sp., 15 Apr.
hyphae, lumen visible; protruding hyphae not seen. Peristomal 2012, J.C. Zamora, L. Papinutti & G. Rolón, holotype MA-Fungi 83764!,
hyphae 2.0 – 6.0 µm wide, light brown, aseptate, mostly un- isotypes in AH 45204! and UPS!
branched, thick-walled (1.0–2.5 µm thick), lumen visible or not, Etymology. The species is dedicated to Dr. L. Papinutti (†), expert on
often sinuous, narrowing at base and apex, tips mostly acute, Argentinian earthstars, who greatly helped the first author during his stay in
a few rounded. Mesoperidium present on the endoperidium Argentina and collected all the studied specimens with him.
and pseudoparenchymatous layer surfaces, consisting of
small, bipyramidal crystals of COD, the largest 6 –13 µm diam, Diagnosis — Exoperidium 12–24 mm diam in horizontal posi-
intermixed with abundant 1.0–3.0 µm wide, hyaline, branched, tion, arched, divided in 6–11 non-hygrometric rays. Endoperidial
thin-walled, clamped hyphae. Pseudoparenchymatous layer of body 3.5–6.5 mm diam, mostly dark brown to blackish. Columel
thin-walled (≤ 1 µm thick), hyaline to yellowish cells, variable in la with a more or less rounded apex. Mesoperidium very abun-
shape and size, about 10 – 65 × 8 – 50 µm diam. Fibrous layer dant, more or less powdery. Largest mesoperidial crystals of
with 1.5 – 4.5 µm wide, hyaline to pale yellowish, aseptate, COD on the endoperidial surface 6.0–12 µm diam, bipyramidal.
straight or slightly sinuous, intertwined, mostly unbranched, Peristome sulcate, distinctly delimited or not, with 12–20 folds.
comparatively thick-walled (0.7– 2.0 µm thick) hyphae, lumen Stalk 0.5–1.3 mm high, normally greyish brown to brown. Basi
visible. Mycelial layer double-layered; inner layer consisting of diospores globose, 4.0–5.5 µm diam, with 0.3–0.6 µm high
1.0–3.0 µm wide, strongly glued together, more or less hyaline, warts, ornamentation verrucose to irregularly pilate. Broadest
branched, thin-walled and clamped hyphae; outer layer with capillitial hyphae 4.5–6.0 µm diam. Pseudoparenchymatous
1.0–3.5(– 4.5) µm wide, hyaline to pale yellowish, aseptate, layer of thin-walled (mostly ≤ 1 µm thick) cells.
rarely branched, comparatively more or less thick-walled (0.5– Macroscopic characteristics — Unexpanded basidiomata
2.0 µm thick) hyphae, lumen very narrow and difficult to see. about 5 mm diam, subglobose and with a rounded apex. Exo
Rhizomorphs with rose-like aggregates of bipyramidal crystals. peridium splitting in 6–11 more or less equal to unequal rays,
7–21 mm diam apparently, 12 – 24 mm diam when forced in whitish to pale-cream coloured. Pseudoparenchymatous layer
horizontal position, arched, often more or less hygrometric. pale cream to greyish cream, not or only superficially cracked,
Mycelial layer thin, whitish to pale cream, strongly intermixed < 0.5 mm thick in dry state, about 1 mm thick when fresh, rather
with debris from the substrate, more or less strongly adhered persistent. Endoperidial body subglobose to ellipsoid, rarely
to the fibrous layer but sometimes peeling-off in some parts. Fi irregular, 3.5–6.5 mm diam, dark brown to blackish; endoperi
brous layer papyraceous to slightly coriaceous when denuded, dial surface glabrous or almost so. Mesoperidium as a rather
Fig. 13   Morphological characters in Geastrum subsect. Sulcostomata pro parte (G. striatum s.l. and G. glaucescens group). a. G. glaucescens (MA-Fungi
83762); b. G. parvistriatum (Zamora 580); c. G. papinuttii (holotype); d. G. striatum (Zamora 242); e. G. aff. striatum (AH 18521). — a1, b1, c1, d1, e1. Basidio
mata habit; a2, b2. median section of the endoperidial body showing the columella with a pointed and persistent apex (a2) or a more rounded and evanescent
apex (b2); c2, d2, e2. detail of the peristome; c3, d3, e3. detail of the stalk an apophysis; a3, b3, c4, d4, e4. mesoperidial crystals of COD on the endoperidial
surface; a4, b4, c5, d5, e5. basidiospores. — Scale bars: a1, b1, c1, d1, e1 = 5 mm; a2, b2, c2, d2, e2, c3, d3, e3 = 2 mm; a3, b3, c4, d4, e4 = 10 µm; a4,
b4, c5, d5, e5 = 2 µm.
thick, more or less powdery, layer of whitish pruina. Peristome Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm
sulcate, with the same colour as the endoperidial surface, thick) cells.
mostly narrowly conical, 0.5 –1.5 mm high, distinctly or indis- Ecology & Distribution — This species grows in siliceous, cal-
tinctly delimited. Stalk present but often short, 0.5–1.3 mm high, careous and gypsiferous soils, under both coniferous and broad-
greyish brown to brown, sometimes light coloured. Apophysis leaf trees, being known only from the central Iberian Peninsula
present or absent, with the same colour as the endoperidium. (Zamora & Calonge 2007, Jeppson 2013). Those records be
Columella intruding about 1/3 –1/2 into the glebal mass, with long to the ‘Mediterranean forests, woodlands and scrub’ biome
a more or less rounded apex. Mature gleba brownish. of the Paleartic ecozone.
Microscopic characteristics — Basidia not seen. Basidio
Additional specimens examined. Spain, Alcalá de Henares, bajo Pinus
spores globose to subglobose, 4.0 – 5.5 µm diam, brownish to halepensis, suelo calizo, 15 Nov. 2011, B. Zamora & J.C. Zamora, herb.
yellowish brown, with 0.3–0.6 µm high brown warts, ornamenta- Zamora 551; Madrid, Ciudad Universitaria, delante del Instituto Anatómico
tion verrucose to irregularly pilate. Broadest capillitial hyphae Forense, bajo Cupressus arizonica var. bonita, 10 Mar. 2005, J.C. Zamora,
4.5–6.0 µm broad, aseptate, very rarely branched, normally MA-Fungi 68580; ibid., under Cupressus arizonica var. bonita, 11 Nov.
straight, thick-walled (2.0 – 2.5 µm thick), with narrow lumen, 2011, J.C. Zamora, herb. Zamora 578; ibid., under Cupressus arizonica var.
mostly visible; tips acute to rounded; surface often covered with bonita, 19 Oct. 2012, J.C. Zamora, C. Galán & L. Zhang, herb. Zamora 539;
Madrid, Ciudad Universitaria, al lado del Instituto de Cardiología de Madrid,
debris. Endoperidium composed of 2.5 – 6.5 µm wide, pale yel-
bajo Cupressus arizonica var. bonita, 10 Mar. 2005, J.C. Zamora, MA-Fungi
lowish to yellowish brown, aseptate, mostly unbranched, slightly 68583; ibid., bajo Cupressus arizonica var. bonita, 26 May 2005, P. Gancedo
sinuous, strongly intertwined, thick-walled hyphae, lumen vis- & J.C. Zamora, MA-Fungi 68582; ibid., under Cupressus arizonica var. bonita,
ible; protruding hyphae not seen. Peristomal hyphae 2.5–7.5 11 Nov. 2011, J.C. Zamora, herb. Zamora 577; Madrid, Getafe, cerro de los
µm wide, light brown, aseptate, mostly unbranched, thick-walled Ángeles, en pinar, 23 Feb. 2008, L. Penelas, herb. Zamora 285; Madrid, Vil-
(1.0–3.0 µm thick), lumen visible or not, more or less straight to laviciosa de Odón, urbanización Campodón, under Olea europaea, 25 Nov.
sinuous, narrowing at apex, tips mostly acute, some rounded. 2007, B. Zamora, herb. Zamora 272; ibid., under Olea europaea, sandy soil,
12 Nov. 2011, B. Zamora & J.C. Zamora, herb. Zamora 580; Toledo, under
Mesoperidium present on the endoperidium and pseudoparen-
Quercus rotundifolia, sandy, siliceous soil, 12 Dec. 2010, J. De Esteban, J.C.
chymatous layer surfaces, consisting of very small, bipyramidal Zamora & B. Zamora, herb. Zamora 460.
crystals, the largest 6.0 –12 diam, intermixed with abundant
1.0–2.5 µm wide, more or less hyaline, branched, thin-walled,
clamped hyphae. Pseudoparenchymatous layer of thin-walled Geastrum pectinatum Pers., Syn. Meth. Fung.: 132. 1801.
(≤ 1 µm thick), hyaline to pale yellowish cells, variable in shape — Fig. 12c
and size, about 12.5–65 × 9–35 µm. Fibrous layer with 1.5–4.0 Type. Lectotype (designated here, MBT198526): f. 11 of pl. XXXVII in
µm wide, hyaline to very pale yellowish, aseptate, straight or Schmidel (1793). Epitype (designated here, MBT198527) that supports the
slightly sinuous, intertwined, mostly unbranched, comparatively lectotype cited above: Sweden, Uppland, Denmark par., between Bergsbrunna
thick-walled (0.7–1.5 µm thick) hyphae, lumen visible. Mycelial Railway Station and Svedden (near Uppsala), on an abandoned ant-hill in
coniferous wood, 5 Sept. 1946, A. Melderis, Lundell & Nannfeldt, Fungi
layer double-layered; inner layer consisting of 1.5–3.0 µm wide,
Exiccati Suecici 1442, UPS 437414 (F-149330)! (Duplicates (isoepitypes)
strongly glued together, more or less hyaline, branched, thin- of this exiccatum were also distributed in the following herbaria: BMI, C, K,
walled and clamped hyphae; outer layer with 1.0–3.0 µm wide, LE, PC, PRM, S, W).
hyaline to somewhat yellowish, aseptate, rarely branched, com-
paratively more or less thick-walled (0.5–1.5 µm thick) hyphae, = Geastrum calyculatum Fuckel, Jahrb. Nassauischen Vereins Naturk.
37. 1870, ‘Geaster calyculatus’.
lumen very narrow and difficult to perceive; myceliar projections
≡ Geastrum bryantii subsp. calyculatum (Fuckel) G. Winter, Dr. L. Raben-
indistinct from the outer myceliar layer. Rhizomorphs covered horst’s Kryptogamen-Flora 1: 911. 1884, ‘Geaster [bryantii subsp.] calycu
with rose-like aggregates of bipyramidal crystals. latus’. — Type: Unknown. In absence of original specimens, f. 3 of t. 5 in
Ecology & Distribution — Only found growing under Prosopis Fuckel (1870) can be selected as lectotype.
spp. in the ‘Tropical and subtropical grasslands, savannas and
shrublands’ biome of the Neotropical ecozone. Synoptic description (based on Sunhede (1989) and studied spe-
cimens) — Exoperidium 20–135 mm diam in horizontal posi-
Additional specimens examined (paratypes). Argentina, Santiago del tion, with (4–)6–10(–11) non-hygrometric rays. Endoperidial
Estero, Guasayán, Prosopis sp., 15 Apr. 2012, J.C. Zamora, L. Papinutti &
body (4.5–)10–25(–36) mm diam, dark brown. Mesoperidium
G. Rolón, MA-Fungi 86911; ibid., Prosopis sp., 15 Apr. 2012, J.C. Zamora,
L. Papinutti & G. Rolón, MA-Fungi 86912.
very abundant, powdery. Largest mesoperidial crystals of
COD on the endoperidial surface 5–15 µm diam, bipyramidal.
Peristome sulcate, distinctly delimited or not, with 11–35 folds.
Geastrum parvistriatum J.C. Zamora & Calonge, Bol. Soc. Stalk slender to more or less stout, (1.5–)3.5–13(–15) mm high
Micol. Madrid 31: 139. 2007. — Fig. 7, 13b and (1.6–)2.0–6.0 mm wide, mostly light coloured. Base of
Type. Spain, Madrid, Ciudad Universitaria, near Instituto de Cardiología de the endoperidial body from smooth to plicate or conspicuously
Madrid, under Cupressus arizonica var. bonita, 24 Nov. 2006, J.C. Zamora, sulcate. Basidiospores globose, 5.5–7.5(–8.0) µm diam, with
holotype MA-Fungi 69583!, isotypes K(M) 147057! and PC 0092573! Numer- 0.6–1.5 µm high warts, ornamentation baculate-pilate. Broadest
ous paratypes were also cited by Zamora & Calonge (2007). capillitial hyphae (8.0 –)9.0 –13.0(–15.0) µm diam. Pseudo
Synoptic description (based on Zamora & Calonge (2007) and parenchymatous layer of thin-walled (mostly ≤ 1 µm thick) cells.
studied specimens) — Exoperidium (13 –)19 – 47 mm diam in Ecology & Distribution — The vast majority of confirmed
horizontal position, arched, divided in 5 –12 non-hygrometric records of this species came from ‘Boreal forests/taiga’, ‘Tem-
rays. Endoperidial body 5 –16 mm diam, mostly dark brown to perate conifer forests’, and ‘Temperate broadleaf and mixed
blackish. Columella with a rounded, rarely somewhat acute, forests’ biomes of the Paleartic ecozone (Sunhede 1989).
rather deciduous apex. Mesoperidium very abundant, powdery. Additional specimens examined. Belgium, sous Picea, 22 Nov. 1964, V.
Largest mesoperidial crystals of COD on the endoperidial surface Demoulin 2889, MA-Fungi 43295. – Spain, Burgos, Aranda de Duero, 19
5.5–12.5 µm diam, bipyramidal. Peristome sulcate, distinctly Oct. 1982; F. Lanata, MA-Fungi 5745; Burgos, jardines junto a la Cartuja de
delimited or not, with 7–23 folds. Stalk 0.5–2.5(–3.0) mm high, Miraflores, bajo Pinus sylvestris, 15 Nov. 2007, F.D. Calonge & F. Esteve-
Raventós, MA-Fungi 75533; Cuenca, terreno básico, bajo Pinus nigra y
mostly brownish to dark brown, rarely pale coloured. Basidio
Juniperus thurifera, Oct. 1988, G. Moreno & J.L. Manjón, AH 11479; Gerona,
spores globose, (4.5–)5.0–7.0(–7.5) µm diam, with (0.4–)0.5– Cerdaña, en humus de conífera, 23 Sept. 2008, E. Vernís, herb. Zamora 292;
1.0(–1.2) µm high warts, ornamentation verrucose to irregularly Huesca, Valle de Hecho, selva de Oza, camino de Espata, bajo Fagus syl
pilate. Broadest capillitial hyphae (5.0–)5.5–8.0(–8.5) µm diam. vatica, Abies alba y Pinus sylvestris, 11 Oct. 2008, F. Palazón, herb. Zamora
290; Lugo, La Rua, bajo Cupressus arizonica, 12 Oct. 1988, J.M. Ruiz, forests’, ‘Tropical and subtropical grasslands, savannas and
MA-Fungi 21686; Lugo, Monforte de Lemos, 12 Oct. 1991, A. Fernández, shrublands’, ‘Temperate grasslands, savannas and shrublands’,
MA-Fungi 28156. – Sweden, Gotland, Gammelgarn parish, Uppstaig natur-
and ‘Montane grasslands and shrublands’ biomes of the Neartic,
reservat, barrskog (coniferous forest), 27 Sept. 2011, T.N. Kristiansen &
Paleartic, and Indo-Malay ecozones (Berkeley 1839).
P. Marstad, UPS F-560803; Gotland, Gothem parish, Jusarve skog, under Picea
abies and Pinus sylvestris, calcareous soil, 30 Sept. 2011, J.C. Zamora, herb. Additional specimens examined (as G. cf. plicatum, which are included
Zamora 613. provisionally under G. plicatum until specimens from the Indo-Malay ecozone
become available for further studies, see Discussion). Argentina, Buenos
Nomenclatural notes — This species could be traced in Per- Aires, partido Lomas de Zamora, Llavallol, Santa Catalina, Ligustrum luci
soon (1794) as ‘G. multifidum var. α’, a name not validly pu dum implanted forest, 12 May 2012, F. Kuhar, V. Castiglia, E. Grassi & J.C.
blished according to Art. 32.1(b), see also Ex. 4 of Art. 9.5. Zamora, MA-Fungi 83774; Entre Ríos, Colón, Ubajay, marginal forest, 2 June
2012, J. Maller & J.C. Zamora, MA-Fungi 87322; Salta, Yungas, 11 Apr. 2012,
Palmer (1959) designated a neotype from Persoon’s herbarium
L. Papinutti, J.C. Zamora & G. Rolón, MA-Fungi 87323. – Tanzania, Iringa,
(see Sunhede 1989) but, according to Art. 9.19(a) of the Njombe, c. 4 km W of Ibumila on Njombe-Kidugale road, S09:10 E34:35,
Melbourne Code (McNeill et al. 2012), it is to be superseded 1820 m, low Uapaca-Brachystegia woodland, 3 June 1985, B. Pettersson,
because original material exists. As previously noted by Sun- M. Hedrén & S.P. Kibuwa 791, UPS F-09935. – Uruguay, Candones, silv.
hede (1989), f. 11–14 of pl. XXXVII in Schmidel (1793), which Cupress, humus aren. sicc. ill. foresta, 40 m, July 1946, Herter, PL. UR.
is the only original material known so far, can be selected as 1908a, Ex. herb. Hert 61024, S.
lectotype. This is formally done here. Plate XXXVII is composed
by several figures, and f. 11 is the one specifically designated Geastrum striatum DC., Fl. Franc. (DC. & Lamarck), ed. 3, 2:
as lectotype because only f. 11 and 12 represent the species 267. 1805, ‘Geaster striatus’ — Fig. 13d
currently know as G. pectinatum, and in f. 11 the stalk with the
pseudoparenchymatous collar as a ring, typical for all species Type. Lectotype (designated here, MBT198542): f. 19 in Bryant (1782).
of the G. pectinatum group, is well-visible. However, since some = Geastrum bryantii Berk., The English Flora of Sir James Edward Smith
specimens of G. pectinatum s.str. and G. meridionale are not 5, 2: 300. 1836, ‘Geaster bryantii ’. — Type: Unknown. In absence of original
easily distinguished on the basis of morphological characters specimens, f. 19 in Bryant (1782) can be selected as lectotype, in which case
(see Discussion), an epitype should also be designated to the name would become a homotypic synonym of G. striatum DC.
= Geastrum orientale Hazsl., Grevillea 6, 39: 108. 1878, ‘Geaster ori
unambiguously set the taxonomic concept of this taxon, in
entalis’. — Type: Unknown. In absence of original specimens, f. 12 –15 of
order to prevent future discussions about its interpretation, and pl. 98 in Hazslinszky (1878) can be selected as lectotype.
preserve the current usage of the name. We prefer to select = Geastrum bryantii subsp. kunzei G. Winter, Dr. L. Rabenhorst’s Krypto
a sequenced, not very old specimen, to provide not only mor- gamen-Flora 1: 911. 1884, ‘Geaster ’. — Type: Unknown.
phological but also molecular phylogenetic data, dispelling any = Geastrum striatum f. rufidum V.J. Staněk in Pilát, Flora ČSR B-1:
uncertainty about its identity. The designated epitype consists 461, 787. 1958. — Type: Czech Republic, Louny, 1 Nov. 1952, M. Křížová,
of three well-preserved, mature fruitbodies in agreement with holotype PRM.
?= Geastrum bryantii var. minor Berk. in Massee, Brit. Fungus-Fl. 1: 37.
Persoon’s (1801) description and with the general concept of
1892, ‘Geaster ’. — Type: Unknown. Besides, Berkeley (in Massee 1892) did
this species, excluding the Macaronesian and Mediterranean not provide bibliographic references or figures.
specimens assignable to G. meridionale. This collection is
part of the Fungi Exiccati Suecici, so duplicates are present in Synoptic description (based on Sunhede (1989) and studied spe-
different herbaria, easing further revisions. The apophysis of cimens)  —  Exoperidium 21–85 mm diam in horizontal posi-
the basidiomata from the epitype varies from smooth to clearly tion, arched, with (4–)6–10(–12) non-hygrometric rays. Endo
sulcate, so it represents part of the intraspecific variation of the peridial body 5.5–26 mm diam, mostly brown to black. Meso
species. In addition, basidiomata were found on an abandoned peridium very abundant, powdery. Largest mesoperidial crystals
ant-hill in coniferous wood, which is one of the preferred habitats of COD on the endoperidial surface 5–15 µm diam, bipyramidal.
of G. pectinatum (Sunhede 1989). Geastrum calyculatum is Apophysis strongly developed, solid, ring-like, with a sharp and
regarded as a synonym based on the chorological, ecological, acute edge. Peristome sulcate, more or less distinctly delimited
and morphological data from the protologue (Fuckel 1870), or not, with 19–36 folds. Stalk (1.0–)2.0–9.0 mm high, whitish to
which fit well within the intraspecific variation of G. pectinatum brownish. Basidiospores globose, 5.0–6.0 µm diam, with 0.5–
observed during our study. 1.0 µm high warts, ornamentation verrucose to irregularly pilate.
Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm
thick) cells.
Geastrum plicatum Berk., Ann. Mag. Nat. Hist. 3: 399. 1839,
Ecology & Distribution — Commonly growing in anthropized
‘Geaster plicatus’. — Fig. 12d
environments, in humic soils often under coniferous, but also broad-
Type. Lectotype (selected by Palmer 1959): India, Madras, Dr. Wight (ex leaf trees. Mainly distributed in ‘Boreal forests/taiga’, ‘Temperate
herb. Hooker), K(M) 180401! conifer forests’, ‘Temperate broadleaf and mixed forests’, and
Synoptic description — Exoperidium 38 – 62 mm diam in hori- ‘Mediterranean forests, woodlands and scrub’ biomes of the
zontal position, with 6 – 9 non-hygrometric rays. Endoperidial Neartic and Paleartic ecozones (Sunhede 1989).
body 10 –18 mm diam, pale to dark brown. Mesoperidium Additional specimens examined. Spain, Madrid, Ciudad Universitaria, near
very abundant, more or less powdery. Largest mesoperidial Instituto de Cardiología de Madrid, under Cupressus arizonica var. bonita,
crystals of COD on the endoperidial surface 7– 20 (–32) µm 8 Oct. 2007, J.C. Zamora, herb. Zamora 242; Madrid, Ciudad Universitaria,
E.T.S.I. de Montes, bajo Pinus halepensis, 9 Jan. 2007, J.C. Zamora, herb.
diam, bipyramidal. Peristome sulcate, distinctly delimited or
Zamora 212; ibid., bajo Cupressus sp., Mahonia aquifolium, Pinus sp. y otras
not, with 16 – 30 folds. Stalk often very slender, 3.5 – 6 mm high plantas, 9 Oct. 2007, J.C. Zamora, herb. Zamora 250; ibid., under Pinus ha
and 0.7–2.5 mm wide, light or dark coloured. Base of the endo lepensis, 17 Oct. 2007, J.C. Zamora, herb. Zamora 251; ibid., bajo Cupressus
peridial body plicate to strongly sulcate. Basidiospores globose, sp., 18 Oct. 2012, J.C. Zamora, L. Zhang & C. Galán, herb. Zamora 540; ibid.,
5.0–6.5 µm diam, with 0.7–1.2 µm high warts, baculate-pilate bajo Pinus halepensis, 18 Oct. 2012, C. Galán, L. Zhang & J.C. Zamora, herb.
ornamentation. Broadest capillitial hyphae 6.0 – 8.0 µm diam. Zamora 541; Valladolid, Villanueva de Duero, encinar de Quercus rotundifo
Pseudoparenchymatous layer of thin-walled (mostly ≤ 1 µm lia, sombrío, húmedo y arenoso, con abundante y gruesa capa de humus,
2 Nov. 2007, G. Martínez & A. García, herb. Zamora 257. – Sweden, Uppland,
thick) cells.
Uppsala, humus of Abies sp., 23 Oct. 2011, J.C. Zamora, MA-Fungi 86672.
Ecology & Distribution — In the broad sense followed in the
present study, this species grows in humic soils. The studied Nomenclatural notes — In agreement with Sunhede (1989),
specimens come from ‘Tropical and subtropical moist broadleaf the names G. bryantii and G. orientale are considered syno-
nyms due to data from the protologues, as well as G. bryantii DISCUSSION

subsp. kunzei. Geastrum striatum f. rufidum is also reduced to
a heterotypic synonym since the presence of reddish tints in An unexpected diversity unravelled by molecular and
the exoperidium is part of the natural intraspecific variation of morphological data
G. striatum. Geastrum bryantii var. minor is only tentatively In the recent taxonomic circumscription of the genus Geastrum
treated as a heterotypic synonym because the data of the proposed by Zamora et al. (2014), Geastrum sect. Geastrum
protologue are vague, so it is not clear which taxon was de- comprised about 17–18 well-defined phylogenetic lineages, but
scribed due to the existence of an European morphologically only 10 species names, i.e., G. arenarium, G. coronatum, G. glau-
close species (G. parvistriatum) that often produces smaller cescens, G. minimum, G. ovalisporum, G. parvistriatum, G. pec-
basidiomata. Finally, Sunhede (1989) noted that f. 19 in Bryant tinatum, G. quadrifidum, G. smithii, and G. striatum, are more
(1782) could be selected as lectotype for G. striatum, but he did or less broadly known and accepted or used in recent studies
not finally lectotypify the species, which is formally done here. (Bates 2004, Fazolino 2009, Jeppson et al. 2013, Kuhar et
al. 2013). In the present study 27 well-defined phylogenetic
Geastrum aff. striatum DC. — Fig. 13e lineages are considered as potential species, which means
almost three times the number of the ‘classical species names’
Synoptic description — Exoperidium 28 – 29 mm diam in hori- previously mentioned. Despite not all specimens could be un
zontal position, more or less arched, with 8–9 non-hygrometric equivocally assigned to a species name, we have been able to
rays. Endoperidial body 9.0–9.5 mm diam, blackish. Mesoperi name 23 species, seven of them proposed as new (G. austro
dium very abundant, powdery. Largest mesoperidial crystals minimum, G. benitoi, G. britannicum, G. kuharii, G. meridionale,
of COD on the endoperidial surface about 12 –13 µm diam, and G. thanatophilum), plus another one (G. leptospermum)
bipyramidal. Apophysis well-developed, solid, ring-like, with recognized by morphological features but not included in
a rounded edge. Peristome sulcate, more or less distinctly molecular analyses, because only the old and scanty mate-
delimited, with 14 – 24 folds. Stalk 1.6 – 2.0 mm high, whitish rial of the type specimen was available. The four subsections
to brownish. Basidiospores globose, 4.5 – 5.0 µm diam, with already defined by Zamora et al. (2014) are retrieved here and
0.4–0.8 µm high warts, ornamentation verrucose to irregularly confirmed as well-defined by both morphological and molecular
pilate. Pseudoparenchymatous layer of thin-walled (mostly data. Geastrum subsect. Hungarica is newly proposed for ac-
≤ 1 µm thick) cells. commodating the morphologically remarkable G. hungaricum
Ecology & Distribution — Insufficiently known; the single (see Taxonomy).
studied collection was found in the ‘Tropical and subtropical
As noted in the results, the single specimen TNS TKG-GE-
dry broadleaf forests’ biome of the northernmost part of the
91002, whose molecular data were collected from GenBank,
Neotropic ecozone.
showed a phylogenetic position intermediate between G. sub-
Additional specimens examined. Mexico, Baja California, sierra La La- sect. Hungarica and G. subsect. Quadrifida. In the GenBank
guna, bajo Lysiloma candida y Pachycereus pringlei, 8 Nov. 1984, C. Ochoa database this specimen is available under the name ‘G. quadri
AH 18521 (duplo BCMEX 1755). fidum’, so one may be tempted to directly include it within G. sub-
sect. Quadrifida, a decision that would not be against our phy-
Geastrum tenuipes Berk., London J. Bot. 7: 576. 1848, logenetic reconstructions (Fig. 1). Unfortunately, this specimen
‘Geaster ’ — Fig. 12e was not available to us for morphological study. Therefore,
making such taxonomic decision without morphological data is
≡ Geastrum pectinatum var. tenuipes (Berk.) Cleland & Cheel, J. Proc. Roy.
Soc. New S. Wales 49: 227. 1915. — Type: Lectotype (selected by Palmer
discouraged, because this specimen may show morphological
1959): Australia, Tasmania, Gunn 1778 (ex herb. M.J. Berkeley), K(M) characters so strongly deviating from other species included in
180399! G. subsect. Quadrifida, for deserving a new subsection. In fact,
as an example, G. britannicum sequences were also available
Synoptic description — Exoperidium 25 – 52 mm diam in hori
as G. quadrifidum in GenBank, but the species is best included
zontal position, with 6 –11 non-hygrometric rays. Endoperidial
in G. subsect. Arenaria (Fig. 1). For these reasons, the speci-
body 8 –17 mm diam, greyish to dark brownish grey. Mesoperi
men TNS TKG-GE-91002 is left taxonomically unplaced.
dium very abundant, slightly powdery or not. Largest meso
peridial crystals of COD on the endoperidial surface (20–)25–55 Due to the complexity of the entire sect. Geastrum, as a result
µm diam, bipyramidal. Peristome sulcate, often distinctly de- of the number of taxa involved and the numerous taxonomic
limited, with 14 – 27 folds. Stalk very slender, 3.0 –7.0 mm high and nomenclatural problems found during this study, comments
and 0.7–1.5 mm wide, light or dark coloured. Base of the endo- and possible solutions proposed for each subsection and spe-
peridial body smooth to plicate or clearly sulcate. Basidiospores cies are separately addressed below.
globose, 5.5 –7.5(– 8.5) µm diam, with 0.7–1.3(–1.5) µm high
warts, ornamentation baculate-pilate. Broadest capillitial hyphae Subsect. Arenaria
6.0–9.5 µm diam. Pseudoparenchymatous layer of thin-walled This subsection is mainly characterized by the shape and size
(mostly ≤ 1 µm thick) cells. of the basidiospores, as well as by the often less marked or-
Ecology & Distribution — Growing on natural and anthropized namentation in comparison with the remaining ones (Zamora
environments, records come from the ‘Temperate broadleaf and et al. 2014). Five species are included within this group in the
mixed forests’ biome of the Australasian ecozone (Berkeley present treatment, of which G. leptospermum is included by
1848, Cunningham 1944). morphological data only.
Additional specimens examined. Australia, Australian Capital Territory, Geastrum ovalisporum, described by Calonge et al. (2000),
Canberra, suburb of Macquarie, S35°15' E149°04', 600 m, suburban garden, is probably the most clearly differentiated species, being the
in dense leaf litter, 7 Nov. 2004, H. Lepp 4603, CANB 745974; Australian
ovoid, small and inconspicuously ornamented basidiospores
Capital Territory, Canberra, Mauldon Street, Chifley, S35°21'20" E149°04'35",
635 m, suburban garden, among leaf and woodchip in garden beds, 5 July
the most distinctive morphological character (Fig. 9e5). The
2008, J. Palmer 760, CANB 775658; Australian Capital Territory, Kaleen, studied specimens are morphologically rather homogeneous;
Canberra, S35°13' E149°06', in a suburban garden, on ground, May 2006, the dark endoperidial body and the whitish, often longer than
C. Evans 9, CANB 738350. broad stalk seem to be reliable characteristics as well.
The South European specimens of the so-called G. arenarium Geastrum leptospermum is one of the smallest earthstars, with
form a clear and well-supported clade according to our phylo- a distinctive combination of morphological characteristics.
genetic analyses (Fig. 1), and deviate from the American taxon Together with G. britannicum and G. quadrifidum, this is the only
which we regarded as G. arenarium s.str. The mesoperidial other fornicate species in G. sect. Geastrum, with basidiospores
crystals of these South European samples are often in the somewhat resembling those of G. britannicum, although smaller
form of bipyramidal prisms with very conspicuous faces even and less ornamented. It also differs by the much smaller basidi-
under the light microscope, mixed with some bipyramids, while omata, very short stalks, absence of big mesoperidial crystal
in the American samples such prisms with conspicuous faces aggregates, and lighter coloured glebal mass (Atkinson 1903,
have not been seen. The disparate distributional and ecological Coker & Couch 1928, Sunhede 1989). The striking ecology
ranges contribute to support the independence of two species. is very different as well, because G. leptospermum grows on
Then, the South European samples are separated as a new mossy bark of living trees instead of growing on soil (Atkinson
species, which is described in the taxonomic part as G. benitoi. 1903, Coker & Couch 1928). Interestingly, we have observed
Although the morphological comparisons of basidiospore size many prism-shaped crystals probably of COM on the rhizo-
and ornamentation height also revealed significant differences, morph surface, forming stellate aggregates (Fig. 2f), in addi-
such differences are of the order of tenths of micrometers and, tion to some bipyramidal crystals of COD. Such crystals were
especially in the case of the ornamentation height, a practi- previously known only in species of G. sect. Myceliostroma,
cal separation is not easy. Therefore, it can be only said that in which they do not form stellate aggregates but are covering
basidiospores in G. arenarium tend to be somewhat smaller cystidioid cells (Zamora et al. 2013). The presence of crystal
and less ornamented than in G. benitoi. Fortunately, and as morphotypes assignable to COM in G. sect. Geastrum is not
noted in the results, G. benitoi is additionally distinguished uncommon, since horn-like crystals have been recorded for
because of its pseudoparenchymatous cells with thin walls, G. ovalisporum, G. subsect. Sulcostomata, and G. coronatum,
while the American G. arenarium specimens have more or less being the main type in the last species (Zamora et al. 2013).
thick walls. According to Sunhede (1989), the thickness of the
pseudoparenchymatous cell wall is related to the hygrometric Subsect. Geastrum
behaviour of the exoperidial rays, and therefore only G. are The subsect. Geastrum, which includes the type of the genus,
narium can be considered as truly hygrometric. In fact, G. are G. coronatum, groups rather stout species with complex basi
narium s.str. is the only species in G. sect. Geastrum, together diospore ornamentation, unique in the genus, made up by big
with G. hungaricum, that has a truly hygrometric exoperidium. processes covered by minute rounded warts (Zamora et al.
Nevertheless, G. hungaricum is clearly distinguished by its 2014), although transitions with subsect. Quadrifida may be
much bigger basidiospores, pseudoparenchymatous cells with observed. The mesoperidium is not powdery, and mesoperidial
thicker walls, more strongly hygrometric exoperidium, mycelial generative hyphae tend to dominate over crystals, which are
layer peeling off very easily, and different crystalline matter of sometimes absent.
the mesoperidium, which is composed almost exclusively by
crystalline aggregates of COM (Fig. 9f3). It should be noted Geastrum smithii is readily distinguished by its regularly sulcate
that the South American specimens of G. arenarium often and sharply delimited, flat peristome. However, specimens shar-
tend to have pseudoparenchymatous cells with slightly thicker ing this morphology and included in the present study are not
walls in comparison with the North American specimens, while completely homogeneous, and probably they represent more
the basidiospores are almost identical. Since few herbarium than one taxon. While the Neartic and Paleartic specimens are
specimens have been studied, we do not know if such subtle morphologically very close, the Australasian specimen differs
differences are consistent, and thus we prefer to treat both mainly by the slightly less stout stalk and the bigger basidio
groups as a single species. All in all, it will not be surprising spores (Fig. 10d3, d5). In addition, mesoperidial crystals are
that future studies allow the distinction of additional species in particularly big in the Australasian specimen, but too few basidio-
this complex. mata have been studied to draw conclusions. The phylogenetic
analyses also show several differences in sequence data (Fig. 1).
Geastrum britannicum is superficially close to G. quadrifidum For these reasons we cannot discard that this specimen may
(G. subsect. Quadrifida) by macromorphological traits, but is belong to a close, but actually different, perhaps undescribed
better included within G. subsect. Arenaria due to both molecular species. Further specimens and molecular data are needed to
data and basidiospore features. In fact, there are several mor- solve the taxonomy of G. smithii s.l.
phological characteristics that can be used to separate G. bri-
tannicum from G. quadrifidum. In G. britannicum the largest On the other hand, the G. coronatum general morphology is
crystals that cover the endoperidial surface are comparatively shared with G. thanatophilum. The true G. coronatum is a rather
big (63 – 80 µm diam) and they are usually grouped and fused common species in Europe, which can be distinguished by the
forming rounded, 60 –120 µm diam aggregates, thus giving a robust and mostly medium-sized basidiomata, the fibrillose peri-
grainy appearance to the endoperidial body, while in G. quadrifi stome, dark stalk, and glabrous endoperidial surface, covered
dum the endoperidium is covered by smaller ((10–)15–50(–60) with a ‘mealy’ mesoperidium formed by abundant but rather
µm), not aggregated, bipyramidal crystals, which confer a indistinct generative hyphae (Fig. 10a; Sunhede 1989). The
pruinose appearance to the endoperidial body. Morphologi- rhizomorph crystals are also rather unusual; they are horn-like
cal analyses also identified that basidiomata size tends to be formations of COM often grouped into arachnoid aggregates
slightly larger in G. britannicum, but too few basidiomata were (Fig. 2e; Zamora et al. 2013).
included in the analyses to consider this a reliable character. Finally, G. thanatophilum is readily distinguished from G. coro-
The same applies to the stalk length, which seems to be slightly natum on account of rhizomorph crystals, mesoperidial features,
longer in G. britannicum. Another macromorphological trait to and capillitial diameter, although old basidiomata of both spe-
distinguish both species is the often inconspicuously folded peri- cies may be easily confused. Then, peristomal characteristics
stome in G. britannicum, which is purely fibrillose in G. quadri may be of some utility when G. thanatophilum has a finely sul-
fidum. In addition, G. quadrifidum has globose, (4.5 –)5.0–6.0 cate peristome, because in G. coronatum it is always fibrillose.
µm diam basidiospores, with very conspicuous, up to 0.8 µm In addition, G. thanatophilum is, on average, a rather smaller
high warts. In contrast, G. britannicum has globose to slightly species (Fig. 4d–f). Geastrum kuharii is another morphologi-
ovoid, (3.5 –) 4.0 – 4.5(– 5.0) µm diam basidiospores, with less cally close species, which belongs to G. subsect. Quadrifida,
conspicuous, up to 0.5(– 0.6) µm high warts. and it is distinguished by the smaller basidiospores, with a less
complex ornamentation, as well as by the common presence and less ornamented basidiospores, and bigger mesoperidial
of crystalline aggregates of COM in the mesoperidium (Fig. 2c crystals (Fig. 3d–f), as previously discussed.
and see below). While G. kuharii and G. quadrifidum are clearly defined spe-
cies, the remaining taxa of this group have been considered
Subsect. Hungarica conspecific and treated under the collective name ‘G. minimum’
This subsection is currently formed by a single species, G. hun- by many authors. In the present treatment, this ‘G. minimum
garicum. This species shares all of the basic traits of G. sect. group’ of putatively cryptic species includes G. austrominimum,
Geastrum as firstly noted by Zamora et al. (2014), except the G. calceum s.l., G. granulosum, and G. marginatum. Thus,
presence of stalk, and is clearly clustered within this section considering that the name G. minimum has been largely applied
in the present phylogenetic analyses (Fig. 1), but cannot be to several species, that the protologue is not enough detailed
properly included in any of the previously defined subsections. to know which species was described by Schweinitz (1822),
Its phylogenetic position is close to G. subsect. Quadrifida, but and that it is difficult to ascertain if the old specimen marked
it posses a remarkable morphology, with a sessile endoperidial as ‘type’ in Kew is really a part of Schweinitz’s original material
body, a strongly hygrometric exoperidium and clearly thick- (Sunhede 1989), we prefer to avoid the use of that name, re-
walled pseudoparenchymatous cells, rather big basidiospores, garding it as nomen ambiguum and dubium until unambiguous
and mesoperidial crystalline matter formed mainly by crystal- original material of this taxon were found. For these reasons,
line aggregates of COM. This combination of morphological comments on the taxonomy of this group are needed to make
characters led us to separate it in a new subsection proposed a proper distinction of the different taxa. Nevertheless, these
here as G. subsect. Hungarica. In relation with the whole genus, four taxa can be distinguished rather well through morphomet-
G. hungaricum can be seen as a morphologically isolated spe- ric analyses (Fig. 5, Table 2), so they are better referred to as
cies that has been considered related to G. floriforme by Staněk pseudo-cryptic species (Mann & Evans 2007, Medina et al.
(1958). However, both species clearly differ from each other, 2012). Nomenclatural notes on certain species are provided
e.g., on the surface of the endoperidium, which is an important in the taxonomic part.
feature for distinguishing several taxa (Sunhede 1989) and for To begin with, all the studied Australasian specimens of ‘G. mini
the infrageneric classification of the genus Geastrum (Zamora et mum s.l.’ are grouped in a well-defined clade (Fig. 1). Although
al. 2014). For that reason, it would have been difficult to include somewhat variable and close to G. marginatum and G. granulo
G. hungaricum in G. sect. Papillata, whose type is G. flori- sum, those specimens are morphologically distinguished by the
forme. Geastrum corollinum is another morphologically close combination of more or less dark coloured stalks, the often rather
species but the mycelial layer does not encrust debris and the big bipyramidal crystals of COD, and particularly by the big, yel-
basidiospore ornamentation is rather different (Sunhede 1989). lowish crystalline aggregates of COM on the endoperidial sur-
In fact, species in G. sect. Corollina as defined by Zamora et face (Fig. 6e, 11a4). Morphometric analyses show that G. mar-
al. (2014), whose type is G. corollinum, mostly show a rather ginatum can be distinguished mainly by the smaller crystals of
regular pilate-baculate ornamentation, in contrast to the irregu- COD and crystalline aggregates of COM on the endoperidial
larly verrucose ornamentation of G. hungaricum (see Fig. 9f4 surface (Fig. 6d, e). In G. granulosum the yellowish crystalline
and Sunhede 1989: 232), which is much closer to that of
aggregates of COM are almost absent or are much smaller,
members of sect. Geastrum, and particularly of G. minimum
although exceptions can be found (Fig. 6e). Fortunately, G. gra-
s.l. (see Fig. 11a5–e5 and Sunhede 1989: 265). The crystalline
nulosum basidiomata show, in addition, light coloured stalks as
aggregates of COM present on the endoperidial surface are
a distinctive qualitative character (Fig. 11d3). Rarely, basidioma-
another important morphological character that places G. hun-
ta of Australasian specimens may show dominant and very big
garicum closer to G. subsect. Quadrifida than to G. sect. Papil
mesoperidial crystals of COD, and then the DFA identified these
lata or G. sect. Corollina.
fruitbodies as belonging to G. granulosum (Fig. 5a). These
basidiomata belong to the voucher MEL 2292062, which is one
Subsect. Quadrifida
of the specimens included in the molecular analyses, nesting
This subsection is a morphologically rather homogeneous group with the remaining Australasian specimens, and not with G. gra-
clustering all the remaining species with a fibrillose peristome. nulosum (Fig. 1). As a result, the Australasian specimens are
It is considerably similar to G. subsect. Arenaria, mainly differ- described here as the new G. austrominimum.
ing in basidiospore shape, size, and ornamentation (Zamora et
Lloyd (1907) described G. calceum (as Geaster calceus) based
al. 2014). In addition, crystalline aggregates of COM are often
on specimens from South Africa. The description is not very
present in some species of this section, but have never been
detailed, but he considered the species as close to G. mini
observed in G. subsect. Arenaria.
mum but being larger and with the endoperidial surface densely
Geastrum kuharii has a remarkable morphology, rather different covered by white granules. Luckily, the iconography (pl. 95)
to other taxa included in G. subsect. Quadrifida, due to the large shows one basidioma with a well-developed, more or less
and stout basidiomata (Fig. 10e1–e3), which strongly remind of dark-coloured stalk, a well-formed apophysis, and a very dense
species in G. subsect. Geastrum, particularly G. thanatophilum crystal covering on the endoperidial surface. This basidioma
as has been commented before. However, basidiospore orna- strongly reminds the specimens included in this study as
mentation is typical of G. subsect. Quadrifida, not as complex G. calceum s.l. However, we have found an unexpected mo-
as in G. subsect. Geastrum, and basidiospores are also smaller, lecular variation between the two sequenced specimens, which
even on the borderline with G. subsect. Arenaria (Fig. 10e5). may indicate that they are different species (Fig. 1). From a
Furthermore, crystalline aggregates of COM are often present morphological point of view, the specimen MA-Fungi 83761
on the endoperidial surface (Fig. 2c) while, as noted above, (G. cf. calceum1) has slightly bigger basidiospores with larger
they have never been seen in species of G. subsect. Geastrum warts and slightly stouter stalks than UFRN-Fungos 723 and
and G. subsect. Arenaria. MA-Fungi 65435 (G. cf. calceum2) (Fig. 11b3–c3, b5–c5). The
Geastrum quadrifidum is a well-known and well-defined spe- ecology and distribution seem to be different, because G. cf.
cies. The fornicate exoperidium, with mostly 4 – 5 rays of con- calceum1 was found in the ‘Tropical and subtropical grasslands,
spicuously rolled margins (Fig. 11f1), is a reliable characteristic savannas and shrublands’ biome, while both specimens of
(Sunhede 1989). The other fornicate species of G. sect. Geas G. cf. calceum2 came from the more humid ‘Tropical and sub-
trum, i.e., G. leptospermum and G. britannicum, have smaller tropical moist broadleaf forests’ biome. Unfortunately, only three
collections of G. calceum s.l. were available for morphological physis. That advises on taking GenBank species names with
study, and among them there was no South African material. caution for determination purposes, as many authors have
Therefore, further data are needed before conclusions on the indicated before for different taxonomic groups (e.g., Nilsson
taxonomy of this species can be drawn. These specimens can et al. 2006, Shen et al. 2013, Sandoval-Sierra et al. 2014). In
be separated from other species of the G. minimum group addition, during our study we have revised one herbarium col-
mainly by the well-marked apophysis (Fig. 11b3 – c3), which is lection from Baja California, Mexico (AH 18521) that was de-
almost ring-like, but also by the ornamentation height and the termined as G. striatum, and previously illustrated by Ochoa &
average basidiomata size (Fig. 6b, c, f). The comparatively Moreno (2006). It consists of two basidiomata that deviate from
slender and long stalks are of some utility, too. Despite the five our concept of G. striatum by having less marked apophyses,
basidiomata included in DFA are classified by such analysis as with rounded edges, and shorter stalks. Basidiospores are also
distinct from the remaining species of the G. minimum group, slightly smaller (4.5–5.0 µm diam). This well-separated sample
it is the only species for which Cohen’s kappa test (Titus et al. is sister to the clade formed by the G. striatum s.str. samples,
1984) was not significant (Table 2), undoubtedly due to the and it very likely represents a different species. Nonetheless,
small sample size. Once more, this data indicate that more having studied only two basidiomata from a single collection,
specimens of this taxon should be studied. we postpone the formal description of this new taxon until more
Fuckel (1860) described G. granulosum (as Geaster granulo specimens are available.
sus) from specimens collected in Germany. According to his Geastrum glaucescens and G. parvistriatum are very close
description, it is a species with fibrillose peristome, and dark, taxa, and their separation is often highly problematic solely
stalked endoperidial body, covered by white granules. Even based on morphology. While well-defined phylogenetically
if the colour of the stalk is not indicated, the white granules, (Fig. 1), the only reliable morphological characteristic found dur-
which the epithet ‘granulosum’ refers to, allow the distinction ing our study is the often rounded columella with a deciduous
of this species from G. marginatum, the only other species of apex in G. parvistriatum (Fig. 13b2), and the normally acute
G. minimum s.l. present in Europe. As indicated in the results, or pointed columella with a rather persistent apex in G. glau-
most basidiomata of G. granulosum were correctly identified cescens (Fig. 13a2). However, a few basidiomata of G. glauces-
by DFA. Basidiomata incorrectly assigned to G. marginatum, cens (particularly the smallest ones) have a more or less
due to particularly small COD crystals, belong to the voucher rounded columella, being almost indistinguishable from most
MICH 72010, and one of the basidiomata incorrectly classified G. parvistriatum basidiomata. In addition, the morphometric
as G. austrominimum, due to particularly big COM crystalline analyses showed that the capillitial hyphae tend to be slightly
aggregates, to the voucher Sunhede 7746. Both specimens broader and the peristome tends to have more folds in G. glau-
were sequenced and they are placed in the same clade as the cescens (Fig. 7d, f), but a wide overlap exists, those differ-
remaining G. granulosum specimens (Fig. 1). ences are not significant, so many specimens cannot be cor-
Geastrum marginatum is the last species of the G. minimum rectly classified by DFA (Fig. 5b). As noted in the results, the
group. Basidiomata typically possess dark stalks (except some only quantitative character that showed significant differences
very small basidiomata), and in morphometric analyses it is dis- was the basidiospore diameter, likely because of the very high
tinguished from G. austrominimum and G. granulosum mainly number of measurements taken, but is obviously useless for
by the sparser and smaller mesoperidial COD crystals and COM taxonomic purposes due to the complete overlap (Fig. 7a). By
crystalline aggregates. Geastrum austrominimum has a very contrast, their disjunct distribution is highly useful to separate
different distribution as commented before, and G. granulosum them, being G. glaucescens a South American and, up to
basidiomata showed light coloured stalks. The few basidiomata now Argentinian species, while G. parvistriatum is a South
incorrectly classified by DFA due to particularly big crystalline European, to date Spanish species. Argentinian records of
elements on the endoperidial surface, showed clearly dark G. parvistriatum (Kuhar et al. 2013) should be referred to as
stalks, which prevents confusion with G. granulosum. When de- G. glaucescens. Taking all these data into account, these two
scribing the species, Vittadini (1842) included the following species can be considered semi-cryptic (Vondrák et al. 2009),
locotypic indication: “In locis aridis arenosis una cum Geastre due to slight differences in morphology and clear dissimilarities
Schmideli circa Papiam frequens. Autumno”. Geastrum mar in ecology and distribution, but not strictly cryptic in the senses
ginatum as conceived in the present study often grows on of Sáez & Lozano (2005) and Mann & Evans (2007).
sandy, siliceous, lime-free to slightly calcareous soils, while Geastrum papinuttii also belongs to the G. glaucescens -G. parvi-
G. granulosum seems to be restricted to, or at least clearly pre- striatum group, but it can be distinguished from both of them
fers, markedly calcareous soils. thanks to the smaller, less ornamented, and paler basidio
spores, the thinner capillitial hyphae, and the smaller and more
Subsect. Sulcostomata delicate basidiomata (Fig. 7a–e). It is likely that some records
This subsection is well-characterized by the sulcate peristome of G. glaucescens mentioned by Kuhar et al. (2013) belong to
and the very well-developed mesoperidium, as a powdery layer this new species.
of very small crystals and generative hyphae on both the endo- Since Palmer’s (1959) revision of the G. plicatum and G. tenui
peridial and pseudoparenchymatous layer surfaces (Zamora et pes type specimens, who considered them as synonyms of
al. 2014). This group is surprisingly diverse and, although only G. pectinatum, this last taxon has been treated as a single and
G. pectinatum and G. striatum are normally taken into consi well-defined species by morphology in all recent studies (e.g.,
deration (plus G. parvistriatum in the most recent treatments), Sunhede 1989, Calonge 1998, Soto & Wright 2000, Bates 2004,
at least 10 taxa can be distinguished according to the results Sarasini 2005, Jeppson et al. 2013). Interestingly, we found a
of the present study. high phylogenetic variation among specimens, strongly related
Geastrum striatum is a well-defined species, and the most to different geographical origins. This suggests a well-defined
striking character is the very well-developed, solid, ring-like geographical structure of several taxa. Therefore, according
apophysis, unique in the genus (Fig. 13d3; Sunhede 1989). to our molecular data, it might be possible to distinguish up
However, two GenBank sequences not generated by us and to five species in this complex, corresponding to four clades
clustering within our clade of G. striatum with strong support from samples sequenced during our study, plus another only
(Fig. 1) were available at the GenBank database under the recovered using GenBank sequences (Fig. 1). This last clade
name G. pectinatum, which never has a solid, ring-like apo- groups exclusively specimens from the Japanese Paleartic
ecozone. It may correspond to G. biplicatum, described from distributed through Europe, from the Temperate to the Boreal
the Bonin Islands by Berkeley & Curtis (1860). Unfortunately, zones, being common in Central and North Europe. Because
only the single basidioma of the lectotype was available for Sunhede’s (1989) description was based on a large number
morphological study. Capillitial hyphae are rather wide, up to of only North European specimens, it is an excellent reference
10.5 µm wide, thus approaching to the often very broad hyphae to the true identity of this species. Surprisingly, two specimens
of G. pectinatum (see below). Basidiospores are comparatively from Japan cluster within the G. pectinatum s.str. clade. More
small, (4.5 –)5.0 – 6.0 µm diam (Fig. 12a5), thus in the range of data are needed to explain such distribution, but a secondary
G. plicatum, but with a less marked ornamentation than any introduction of the species in that country should not be dis-
other species of the G. pectinatum group, with warts hardly carded, as Geastrum specimens are not rarely found in parks
reaching 1 µm high. Mesoperidial crystals of COD are small (up and gardens.
to 15 µm diam; Fig. 12a4) as in the other species of G. subsect. The other European clade groups the Macaronesian and Medi
Sulcostomata except G. tenuipes and sometimes G. plicatum. terranean specimens, which are close to G. pectinatum but
Regrettably, the sequenced specimens that might correspond clearly separated by molecular data (Fig. 1). These specimens
to G. biplicatum due to geographical origin were not available are described in this study as G. meridionale. Although the
to us for morphological study, and thus their identity is only representation of the first two discriminant functions of the DFA
suspected and could not be confirmed. What is more, while may give the false impression that these two taxa are hardly
the sequenced specimens belong to the Paleartic ecozone, the distinguished by the studied morphological characters (Fig. 5c),
type specimen was collected in the Oceanian ecozone, so the the high percentage of correctly classified samples (95 % of
existence of more than one species would not be surprising. G. meridionale and 100 % of G. pectinatum) indicates that both
The other four phylogenetic clades have been morphologically species are more or less well-separated by morphology. The
studied with more detail, and morphometric analyses (Fig. 5, overlap in the score plot is explained because the third discri-
Table 2) indicate a complex of morphologically close, but dis minant function is the one that best distinguishes G. pectinatum
tinguishable, pseudo-cryptic species (Mann & Evans 2007, and G. meridionale. One of the misclassified basidiomata of
Medina et al. 2012). The clade formed by both Paleo- and G. meridionale belongs to the voucher MA-Fungi 20651, which
Neotropical specimens of G. pectinatum s.l. may correspond was included in the phylogenetic analyses and clusters together
to G. plicatum. This taxon was described from India, Madras with the remaining samples of G. meridionale (Fig. 1). While
(Berkeley 1839), and referred to as specimens having long and G. pectinatum has the thickest capillitium hyphae, G. meridio
thin stalks and strongly sulcate apophysis or endoperidial basis, nale has the stoutest stalks of the entire complex (Fig. 8e, 12b3).
in agreement with most of the tropical specimens studied. The In addition, the number of peristome folds of G. meridionale is
basidiospores are one of the smallest of the complex (Fig. 8a, normally much higher, and basidiomata tend to be lighter, espe-
12d5), while the grooves on the apophysis or the base of the cially the endoperidial body, which is mostly cream coloured to
endoperidial body are the most conspicuously marked ones. brownish, and never blackish as is commonly seen in G. pec-
The type specimen of G. plicatum agrees rather well with the tinatum. Nevertheless, dark basidiomata of G. meridionale with
specimens included in our molecular analyses, although the slender stalks and a low number of peristome folds can be very
mesoperidial crystals tend to be larger and basidiospores difficult to distinguish from G. pectinatum, especially when old;
measure 5.5 – 6.5 µm diam, being transitional with G. tenuipes then, molecular data become a useful tool for a more precise
(see below). This specimen was included in the DFA of the classification.
G. pectinatum group (Fig. 5c), clustering with other basidiomata To end, since some extensive areas were not sampled (e.g.,
determined as G. plicatum. However, since no samples from the North America, continental Asia, and Indonesia), other taxa be
Indo-Malay ecozone have been sequenced, the name G. pli- longing to the G. pectinatum complex might be missing. Further
catum is only tentatively assigned to the Neotropical and Afro- worldwide-based sampling would be needed to propose a more
tropical (Ethiopian) specimens. definite taxonomic treatment.
The Australasian specimens of G. pectinatum s.l. are phyloge-
netically well-defined (Fig. 1) and would correspond to the so-
CONCLUSIONS
called G. tenuipes. This species was described from Tasmania
and Australia (Berkeley 1848) based on slender specimens with
The example of fungal species delimitation showed with G. sect.
rather long and thin stalks, and slightly sulcate endoperidial
Geastrum indicates that, even in hardly morphologically dis-
bases. These characteristics match fairly well with the recently
tinguished species, an integrative taxonomy, as a sum of all
collected Australasian specimens revised during this study,
the taxonomic information sources available (i.a., molecular,
although the depth of the basal endoperidial grooves seems
morphological, chorological, and ecological data) may result
variable even within the same collection. The mesoperidial
in taxonomic partitions recognizable at species level.
crystals are constantly bigger than in the remaining G. pectina
tum group samples (Fig. 8c, 12e4), and seem to be a reliable More specifically, in our study, phylogenetic analyses have
characteristic for distinguishing this taxon, although they have been basic to establish a solid and objective taxonomic back-
not been mentioned previously in the literature. Basidiomata bone, and to identify nearly cryptic taxa. Discriminant function
from the type specimen are very old and the mesoperidium is analyses have been successfully used as a tool for assessing
almost lost; as a result, the mesoperidial crystal size cannot morphological boundaries, and identifying morphological char-
be evaluated. Basidiospores are, on average, somewhat small, acters that may be useful for species characterization. Ecologi-
overlapping with G. plicatum, but within the range observed in cal and chorological data contribute to support the taxonomic
recently collected specimens of G. tenuipes. partitions observed.
As previously noted, the European specimens of G. pectinatum
s.l. form two well-defined clades. One of them belongs to G. pec- Acknowledgements   We thank the curators of the studied herbaria for send-
ing specimens on loan, especially to M. Dueñas (MA-Fungi), and all people
tinatum s.str., a morphologically very variable taxon with mostly
who contributed directly providing specimens for our studies. Particularly
slender but sometimes more or less stout stalks, showing also warm thanks are due to people who greatly helped the first author during his
a broad variation concerning the folds of the apophysis (Sun- stays in Sweden (S. Sunhede, S. Ekman, S. Rymman, M. Myrdal, K. Hansen,
hede 1989). Capillitium hyphae are the widest of the complex and I. Olariaga) and Argentina (L. Papinutti (†), F. Kuhar, V. Castiglia,
(Fig. 8b) and seem to be a reliable characteristic. It is widely G. Rolón, E. Grassi, S. Suaza, A. Romero, A. Arambarri (†), and E. García).
We thank Yolanda Ruiz for her technical assistance with the SEM. Isabel Frey-Wyssling A. 1981. Crystallography of the two hydrates of crystalline
M. Liberal, P. Jiménez-Mejías and J.V. Sandoval-Sierra kindly provided calcium oxalate in plants. American Journal of Botany 68, 1: 130 –141.
information on some morphological analyses. Luis A. Parra kindly helped Fuckel L. 1860. Enumeratio Fungorum Nassoviae. Jahrbücher des Nassaui
with comments on some nomenclatural issues. Joaquina M. García-Martín is schen Vereins für Naturkunde 15: 1–123.
acknowledged for revising the English wording. The first author was supported Fuckel L. 1870. Symbolae Mycologicae. Jahrbücher des Nassauischen Ver
by a predoctoral fellowship from the Consejo Superior de Investigaciones eins für Naturkunde 23: 1– 459.
Científicas (Jae-Pre 2010). Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for ba-
sidiomycetes. Application to the identification of mycorrhizae and rusts.
Molecular Ecology 2: 113 –118.
REFERENCES Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor
and analysis program for Windows 95/98/NT. Nucleic Acids Symposium
Atkinson GF. 1903. A new species of Geaster. Botanical Gazette 36: 303 – Series 41: 95 – 98.
306. Hazslinszky FA. 1878. On Geaster orientalis nov. spec. Grevillea 6, 39: 108–
Bartlett MS. 1937. Properties of sufficiency and statistical tests. Proceedings 109.
of the Royal Statistical Society, Series A 160: 268 – 282. Hazslinszky FA. 1883. Bemerkungen zu den deutschen und ungarischen
Bates ST. 2004. Arizona members of the Geastraceae and Lycoperdaceae Geaster-Arten. Verhandlungen des Botanischen Vereins der Provinz Bran
(Basidiomycota, Fungi). Master thesis, Arizona State University, USA. denburg und die angrenzenden Lander 24: 135 –137.
Berkeley MJ. 1839. Descriptions of exotic fungi in the collection of Sir W. J. Henderson A. 2006. Traditional morphometrics in plant systematics and its
Hooker, from memoirs and notes of J.F. Klotzsch, with additions and cor- role in palm systematics. Botanical Journal of the Linnean Society 151:
rections. Annals and Magazine of Natural History, series 1, 3: 375 – 401. 103 –111.
Berkeley MJ. 1848. Decades of fungi. Hooker’s London Journal of Botany Hillis DM, Bull JJ. 1993. An empirical test of bootstrapping as a method for
7: 572–580. assessing confidence in phylogenetic analyses. Systematic Biology 42:
Berkeley MJ, Curtis MA. 1860. Characters of new fungi, collected in the 182–192.
North Pacific Exploring Expedition by Charles Wright. Proceedings of the Hollós L. 1901. Új gasteromyceta fajok magyarországból. Mathematikai és
American Academy of Arts and Sciences 4: 110 –130. természettudományi értesítõ 19: 504 – 512.
Borchsenius F. 2007. FastGap 1.0.8. Software distributed by the author.
Hollós L. 1904. Die Gasteromyceten Ungarns. Weigel, Leipzig, Germany.
Bottomley AM. 1948. Gasteromycetes of South Africa. Bothalia 4: 473 – 810.
Horner HT, Tiffany LH, Knaphus G. 1995. Oak-leaf-litter rhizomorphs from
Brown JM, Hedtke SM, Lemmon AR, et al. 2010. When trees grow too long:
Iowa and Texas: Calcium oxalate producers. Mycologia 87, 1: 34 – 40.
Investigating the causes of highly inaccurate Bayesian branch-length
Jeppson M. 2013. Jordstjärnor. Mykologiska publikationer 6: 1– 228.
estimates. Systematic Biology 59: 145 – 161.
Jeppson M, Nilsson RH, Larsson E. 2013. European earthstars in Geastra
Bryant C. 1782. An historical account of two species of Lycoperdon. London.
ceae (Geastrales, Phallomycetidae) – a systematic approach using mor
Calonge FD. 1998. Gasteromycetes. I. Lycoperdales, Nidulariales, Phal-
phology and molecular sequence data. Systematics and Biodiversity 11,
lales, Sclerodermatales, Tulostomales. Flora Mycologica Iberica 3: 1–271.
4: 437– 465.
Calonge FD, Moreno-Arroyo B, Gómez J. 2000. Aportación al conocimiento
Jiménez-Mejías P, Luceño M, Martín-Bravo S. 2014. Species boundaries
de los Gasteromycetes, Basidiomycotina, de Bolivia (América del Sur).
within the southwest Old World populations of the Carex flava group (Cy-
Geastrum ovalisporum sp. nov. Boletín de la Sociedad Micológica de
peraceae). Systematic Botany 39, 1: 117–131.
Madrid 25: 271–276.
Kaiser HF. 1974. An index of factorial simplicity. Psychometrika 39: 31– 36.
Calonge FD, Zamora JC. 2003. Geastrum arenarium, encontrado en España
Kasuya T, Hosaka K, Uno K, et al. 2012. Phylogenetic placement of Geas-
y nuevo para Europa. Boletín de la Sociedad Micológica de Madrid 27:
trum melanocephalum and polyphyly of Geastrum triplex. Mycoscience
59 – 61.
53, 6: 411– 426.
Carstens BC, Pelletier TA, Reid NM, et al. 2013. How to fail at species
Katoh K, Misawa K, Kuma K, et al. 2002. MAFFT: a novel method for rapid
delimitation. Molecular Ecology 22, 17: 4369 – 4383.
multiple sequence alignment based on fast Fourier transform. Nucleic Acids
Castresana J. 2000. Selection of conserved blocks from multiple alignments
Research 30: 3059 – 3066.
for their use in phylogenetic analysis. Molecular Biology and Evolution 17,
Kretzer A, Bruns TD. 1999. Use of atp6 in fungal phylogenetics: an example
4: 540–552.
from the Boletales. Molecular Phylogenetics and Evolution 13: 483 – 492.
Chajewski M. 2009. rela: Item Analysis Package with Standard Errors. R
Kuhar F, Castiglia V, Papinutti L. 2013 ‘2012’. Geastrum species of the La
package version 4.1.
Rioja province. Mycotaxon 122: 145 –156.
Coker WC, Couch JN. 1928. The Gasteromycetes of the Eastern United
Lewis PO. 2001. A likelihood approach to estimating phylogeny from discrete
States and Canada. The University of Carolina Press, USA.
Cortez VG, Sulzbacher MA, Baseia IG, et al. 2008. Two little known gasteroid morphological character data. Systematic Biology 50, 6: 913– 925.
fungi from Santa Catarina State, southern Brazil. Mycotaxon 106: 297–302. Lloyd CG. 1902. The Geastrae. Bulletin of the Lloyd Library of Botany, Phar
Cubeta MA, Echandi E, Abernethy T, et al. 1991. Characterization of anasto- macy and Materia Medica 2: 1– 43.
mosis groups of binucleate Rhizoctonia species using restriction analysis Lloyd CG. 1907. New notes on the Geasters. Mycological Writings 2: 309 –
of an amplified ribosomal RNA gene. Phytopathology 81: 1395 –1400. 317.
Cunningham GH. 1944 [reprint 1979]. The Gasteromycetes of Australia and Lutzoni F, Kauff F, Cox CJ, et al. 2004. Assembling the Fungal Tree of Life:
New Zealand. Bibliotheca Mycologica 67: 1– 239. progress, classification, and evolution of sucellular traits. American Journal
Darriba D, Taboada GL, Doallo R, et al. 2012. jModelTest 2: more models, of Botany 91, 10: 1446 –1480.
new heuristics and parallel computing. Nature Methods 9, 8: 772. Macbride TH. 1912. Notes on Iowa saprophytes – I. Geaster minimum Schw.
Dayrat B. 2005. Towards integrative taxonomy. Biological Journal of the and its relatives. Mycologia 4, 2: 84 – 87.
Linnean Society 85: 407– 415. Mann DG, Evans KM. 2007. Molecular genetics and the neglected art of dia-
Demoulin V. 1984. Typification of Geastrum Pers. : Pers. and its orthographic tomics. In: Brodie J, Lewis J (eds), Unravelling the algae: the past, present
variant Geaster (Gasteromycetes). Taxon 33, 3: 498 – 501. and future of algal systematics: 232 – 265. CRC Press, USA.
Edwards DL, Knowles LL. 2014. Species detection and individual assignment Marshall DC. 2010. Cryptic failure of partitioned Bayesian phylogenetic
in species delimitation: can integrative data increase efficacy? Proceedings analyses: Lost in the land of long trees. Systematic Biology 59: 108 –117.
of the Royal Society B 281: 20132765. Martín MP, Winka K. 2000. Alternative methods of extracting and amplifying
Ekman S, Blaalid R. 2011. The devil in the details: interactions between the DNA from lichens. Lichenologist 32, 2: 189 –196.
branch-length prior and likelihood model affect node support and branch Massee G. 1892. British Fungus Flora 1. Clowes & Sons, London, United
lengths in the phylogeny of Psoraceae. Systematic Biology 60, 4: 541–561. Kingdom.
Eyndhoven GL. 1937. Geastrologische notizen. Mededelingen van de Neder Matheny PB, Liu YJ, Ammirati JF, et al. 2002. Using RPB sequences to
landse Mycologische Vereeniging 24: 12 –19. improve phylogenetic inference among mushrooms (Inocybe, Agaricales).
Fazolino E. 2009. O gênero Geastrum Pers. (Phallomycetidae, Basidio American Journal of Botany 89, 4: 688 – 698.
mycota) em algumas áreas de Mata Atlântica e Caatinga no Rio Grande McNeill J, Barrie FR, Buck WR, et al. 2012. International Code of Nomen-
do Norte, Brasil. Master thesis. Universidade Federal do Rio Grande do clature for algae, fungi, and plants (Melbourne Code), adopted by the
Norte, Natal, Brazil. Eighteenth International Botanical Congress Melbourne, Australia, July
Felsenstein J. 1985. Confidence limits on phylogenies: An approach using 2011. Regnum Vegetabile 154.
the bootstrap. Evolution 39: 783 –791. Medina R, Lara F, Goffinet B, et al. 2012. Integrative taxonomy successfully
Fisher RA. 1936. The use of multiple measurements in taxonomic problems. resolves pseudo-cryptic complex of disjunct epiphytic moss Orthotrichum
Annals of Eugenics 7: 179 –188. consimile s.l. (Orthotrichaceae). Taxon 61: 1180 –1198.
Nilsson RH, Ryberg M, Kristiansson E, et al. 2006. Taxonomic reliability Shen Y-Y, Chen X, Murphy RW. 2013. Assessing DNA barcoding as a tool
of DNA sequences in public sequence databases: a fungal perspective. for species identification and data quality control. PLoS ONE 8, 2: e57125.
PLoS ONE 1, 1: e59. Simmons MP, Ochoterena H. 2000. Gaps as characters in sequence-based
Nylander JAA, Wilgenbusch JC, Warren DL, et al. 2008. AWTY (are we phylogenetic analyses. Systematic Biology 49: 369 – 381.
there yet?): a system for graphical exploration of MCMC convergence in Šmarda F. 1947. Nový druh hvězdice – Geaster atratus sp. n. (hvězdice
Bayesian phylogenetics. Bioinformatics 24: 581– 583. černavá). Česká Mykologie 1: 71–74.
Ochoa C, Moreno G. 2006. Hongos gasteroides y secotioides de Baja Cali- Smith JA, Blanchette RA, Newcombe G. 2004. Molecular and morphologi-
fornia, México. Boletín de la Sociedad Micológica de Madrid 30: 121–166. cal characterization of the willow rust fungus, Melampsora epitea, from
Olson DM, Dinerstein E, Wikramanayake ED, et al. 2001. Terrestrial eco artic and temperate hosts in North America. Mycologia 96, 6: 1330 –1338.
regions of the world: A new map of life on earth. BioScience 51: 933 – 938. Soto MK, Wright JE. 2000. Taxonomía del género Geastrum (Basidiomycetes,
Palmer JT. 1959. Observations on Gasteromycetes. VIII. Persoon’s speci- Lycoperdales) en la provincia de Buenos Aires, Argentina. Boletín de la
mens of Geastrum pectinatum Pers. and a reassessment of Geastrum Sociedad Argentina de Botánica 34, 3 – 4: 185 – 201.
plicatum Berk. and G. tenuipes Berk. Persoonia 1: 149 –164. Stamatakis A. 2006. RAxML-VI-HPC: Maximum likelihood-based phyloge-
Persoon CH. 1794. Neuer Versuch einer systematischen Einteilung der netic analyses with thousands of taxa and mixed models. Bioinformatics
Schwämme. Neues Magazin für die Botanik 1: 63 –128. 22, 21: 2688 – 2690.
Persoon CH. 1801. Synopsis Methodica Fungorum. Apud Henricum Dietrich, Staněk VJ. 1958. Geastraceae. In: Pilát A (ed), Flora ČSR B1, Gasteromy-
Germany. cetes: 392 – 526, 777–795. Praha.
Pimentel RA. 1979. Morphometrics, the multivariate analysis of biological Stevens J. 1996. Applied multivariate statistics for the behavioral sciences
data. Kendall, Hunt Pub. Co. (3rd ed). Mahwah, NJ, Lawrence Erlbaum.
Quélet L. 1873. Les champignons des Jura et des Vosges, vol. 2. Mémoires Stiller JW, Hall BD. 1997. The origin of red algae: implications for plastid
de la Société d’émulation de Montbéliard 2, 5: 43 – 332. evolution. Proceedings of the National Academy of Sciences, USA 94:
R Development Core Team. 2008. R: A language and environment for sta- 4520 – 4525.
tistical computing. R Foundation for Statistical Computing, Vienna, Austria. Sunhede S. 1986. Notes on the type material of Geastrum arenarium. Wind
http://www.R-project.org. halia 15: 35 – 43.
Rabenhorst L. 1851. Mykologisches I. Botanische Zeitung 9: 625 – 629. Sunhede S. 1989. Geastraceae (Basidiomycotina). Morphology, ecology and
Rambaut A. 2007. Molecular evolution, phylogenetics and epidemiology. systematics with special emphasis on the North European species. Synop
FigTree v1.3. http://tree.bio.ed.ac.uk/software/figtree/. sis Fungorum 1: 1– 534.
Rambaut A, Suchard MA, Xie D, et al. 2013. Tracer v1.5, Available from Swofford DL. 2003. PAUP*: Phylogenetic Analysis Using Parsimony (*and
http://beast.bio.ed.ac.uk/Tracer. other methods), v. 4.0b10. Sinauer Associates, Sunderland, Massachusetts.
Rannala B, Zhu T, Yang Z. 2012. Tail paradox, partial identifiability and Titus K, Mosher JA, Williams BK. 1984. Chance-corrected classification for
influential priors in Bayesian branch length inference. Molecular Biology use in discriminant analysis: Ecological applications. American Midland
and Evolution 29: 325–335. Naturalist 111: 1–7.
Revelle W. 2014. psych: Procedures for Personality and Psychological Valcárcel V, Vargas P. 2010. Quantitative morphology and species delimitation
Research, Northwestern University, Evanston, Illinois, USA, http://CRAN.R- under the general lineage concept: Optimization for Hedera (Araliaceae).
project.org/package=psych. Version 1.4.4. American Journal of Botany 97, 9: 1555 –1573.
Roca-Valiente B. 2013. Phylogenetic study in the Rhizocarpon geographicum Valdecasas AG, Williams D, Wheeler QD. 2008. ‘Integrative taxonomy’ then
group (Lichens, Rhizocarpaceae, Ascomycota). Contrasting analysis of and now: a response to Dayrat (2005). Biological Journal of the Linnean
morphological characters and biogeographic patterns. PhD thesis, Biología Society 93: 211– 216.
Vegetal II, Universidad Complutense de Madrid, Spain. Vilgalys R, Hester M. 1990. Rapid genetic identification and mapping of
Ronquist F, Teslenko M, Mark P van der, et al. 2012. MrBayes 3.2: Efficient enzymatically amplified DNA from several Cryptococcus species. Journal
Bayesian phylogenetic inference and model choice across a large model of Bacteriology 172: 4238 – 4246.
space. Systematic Biology 61, 3: 539 – 542. Vittadini C. 1842. Monographia Lycoperdineorum. Ex Officina Regia, Italy.
Ruiz-Sánchez E, Sosa V. 2010. Delimiting species boundaries within the Vondrák J, Říha P, Arup U, et al. 2009. The taxonomy of the Caloplaca citrina
Neotropical bamboo Otatea (Poacaeae: Bambusoideae) using molecular, group (Teloschistales) in the Black Sea region; with contributions to the
morphological and ecological data. Molecular Phylogenetics and Evolu- cryptic species concept in lichenology. The Lichenologist 41: 571– 604.
tion 54: 344–356. White TJ, Bruns T, Lee S, et al. 1990. Amplification and direct sequencing
Sáez AG, Lozano E. 2005. Body doubles. Nature 433: 111. of fungal ribosomal RNA genes for phylogenetics. In: Innis M, Gelfand J,
Sandoval-Sierra JV, Martín MP, Diéguez-Uribeondo J. 2014. Species Sninsky J, et al., PCR protocols: a guide to methods and applications:
identification in the genus Saprolegnia (Ooomycetes): Defining DNA-based 315 – 322. Academic Press, Orlando, Florida.
molecular operational taxonomic units. Fungal Biology 118: 559 – 578. Zamora JC. 2014. Proposal to conserve the name Geastrum (Basidiomycota:
Sarasini M. 2005. Gasteromiceti Epigei. Associazione Micologica Bresadola, Geastrales) with a conserved type. Taxon 63, 3: 679 – 680.
Italy. Zamora JC, Calonge FD. 2007. Geastrum parvistriatum, una nueva especie
Schaeffer JCh. 1763. Fungorum, qui in Bavaria et Palatinau circa Ratisbonam encontrada en España. Boletín de la Sociedad Micológica de Madrid 31:
nascuntur icones, nativis coloribus expressae 2. Henrici Godofredi Zunkelii, 139 –149.
Ratisbona, Germany. Zamora JC, Calonge FD, Hosaka K, et al. 2014. Systematics of the genus
Schlick-Steiner BC, Steiner FM, Seifert B, et al. 2010. Integrative taxonomy: Geastrum (Fungi: Basidiomycota) revisited. Taxon 63, 3: 477– 497.
a multisource approach to exploring biodiversity. Annual Review of Ento- Zamora JC, Calonge FD, Martín MP. 2013. New sources of taxonomic infor
mology 55: 421–438. mation for earthstars (Geastrum, Geastraceae, Basidiomycota): phenoloxi-
Schmidel DCC. 1793. Icones plantarum et analyses partium aeri incisae dases and rhizomorph crystals. Phytotaxa 132, 1: 1– 20.
atque vivis coloribus indicibus nominum necessariis figuram explicationibus Zhang C, Rannala B, Yang Z. 2012. Robustness of compound Dirichlet priors
et brevibus animadversionibus, Ed. 2. Erlangae. for Bayesian inference of branch lengths. Systematic Biology 61: 779–784.
Schweinitz LD. 1822. Synopsis fungorum Carolinae superioris secundum Zwickl DJ. 2006. Genetic algorithm approaches for the phylogenetic analy-
observations. Schriften der Naturforschenden Gesellschaft zu Leipzig 1: sis of large biological sequence datasets under the maximum likelihood
20 –130. criterion. PhD thesis, The University of Texas at Austin, USA.

Geastrum Sect Geastrum 1422974412556

Uploaded by

Document Informationclick to expand document information

Document Informationclick to expand document information

Copyright:

Available Formats

Geastrum Sect Geastrum 1422974412556

Uploaded by

Document Information

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Geastrum Sect Geastrum 1422974412556

Uploaded by

Copyright:

Available Formats

Persoonia 34, 2015: 130 –165

www.ingentaconnect.com/content/nhn/pimj RESEARCH ARTICLE http://dx.doi.org/10.3767/003158515X687443

Integrative taxonomy reveals an unexpected diversity in

INTRODUCTION It is known that different species delimitation approaches can

© 2015 Naturalis Biodiversity Center & Centraalbureau voor Schimmelcultures

G. arenarium Argentina, La Rioja MA-Fungi 83760 KF988351 KF988471 KF988606 KF988741

G. quadrifidum Sweden, Uppland MA-Fungi 86671 KF988422 KF988550 KF988685 KF988817

G. pectinatum TNS TKG-GE-91201

Geastrum arenarium / Geastrum benitoi

Pseudoparenchymaous cell walls

Geastrum britannicum / Geastrum leptospermum / Geastrum quadrifidum

COD crystal diameter

0.5 1.0 1.5 2.0 2.5 3.0

BRI LEP QUA BRI LEP QUA BRI LEP QUA

Geastrum coronatum / Geastrum kuharii / Geastrum thanatophilum

COD crystal diameter

COR KUH THA COR KUH THA COR KUH THA

COR KUH THA COR KUH THA COR KUH THA

G. minimum group G. glaucescens and G. parvistriatum were only found in basidio­

1 COD signiﬁcantly larger than other species (Fig. 8c). Geastrum

Geastrum section Geastrum

1. Peristome always sulcate; mostly conical to narrowly coni-

1. Peristome often ﬁbrillose, rarely sulcate; mostly flat to broadly

Geastrum minimum group (G. austrominimum / G. calceum / G. granulosum / G. marginatum)

Maximum ornamentation height

COM aggregate diameter

Geastrum glaucescens group (G. glaucescens / G. papinuttii / G. parvistriatum)

GLA PAP PAR GLA PAP PAR GLA PAP PAR

GLA PAP PAR GLA PAP PAR GLA PAP PAR

Geastrum pectinatum group (G. meridionale / G. pectinatum / G. plicatum / G. tenuipes)

COD crystal diameter

Stalk length/stalk width

nyms due to data from the protologues, as well as G. bry­antii DISCUSSION

You might also like

G. minimum group G. glaucescens and G. parvistriatum were only found in basidio

nyms due to data from the protologues, as well as G. bryantii DISCUSSION