Some interesting gasteroid and secotioid fungi from Sonora, Mexico

Abstract

Nine rare species of gasteroid and secotioid fungi from Sonora, Mexico are treated here: Agaricus texensis (= Longula texensis), Araneosa columellata, Calvatia bicolor, C. craniiformis, C. pygmaea, Disciseda hyalothrix, D. verrucosa, Endoptychum arizonicum, and D. stuckertii (= Abstoma stuckertii), which is a new combination. SEM micrographs of all studied taxa are included. Fungal diversity makes the Mexican State of Sonora highly interesting. The vegeta-tion comprises several types, namely microphyllous desert scrub, tropical thorn forest, tropical deciduous forest, oak and oak-pine forest. Many of these fungi tolerate long dry periods, which characterize arid and semi-arid regions (200 – 350 mm / yr) and play a decisive role in the conservation of ecosystems, especially in the recycling of organic matter. The present work focuses on these mushrooms of agaricoid and gastroid habits, traditionally classified in the artificial group Gasteromycetes. Nine species of mushrooms, which are little known world-wide or which occur in a very small area, were identified from samples collected during the last decade in Sonora. These include Endoptychum arizonicum, a gregarious species limited to the desert between Mexico and the USA; Araneosa columellata, recorded for the second time by Esqueda et al. (1998), 60 years after the holotype was registered from Arizona, USA (long, 1941); and Calvatia pygmaea, reported by Fries (1909) from Argentina and Bolivia, and found again in Baja California by Ochoa et al. (1998).

Full text

SOME INTERESTING GASTEROID AND SECOTIOID
FUNGI FROM SONORA, MEXICO
G. MORENO1, M. ESQUEDA2, E. PÉREZ-SILVA3, T. HERRERA3 & A. ALTÉS1
Nine rare species of gasteroid and secotioid fungi from Sonora, Mexico are treated
here: Agaricus texensis (= Longula texensis), Araneosa columellata, Calvatia bicolor,
C. craniiformis, C. pygmaea, Disciseda hyalothrix, D. verrucosa, Endoptychum
arizonicum, and D. stuckertii (= Abstoma stuckertii), which is a new combination.
SEM micrographs of all studied taxa are included.
Fungal diversity makes the Mexican State of Sonora highly interesting. The vegeta-
tion comprises several types, namely microphyllous desert scrub, tropical thorn forest,
tropical deciduous forest, oak and oak-pine forest. Many of these fungi tolerate long
dry periods, which characterize arid and semi-arid regions (200 – 350 mm / yr) and play
a decisive role in the conservation of ecosystems, especially in the recycling of organic
matter. The present work focuses on these mushrooms of agaricoid and gastroid
habits, traditionally classified in the artificial group Gasteromycetes. Nine species of
mushrooms, which are little known world-wide or which occur in a very small area,
were identified from samples collected during the last decade in Sonora. These include
Endoptychum arizonicum, a gregarious species limited to the desert between Mexico
and the USA; Araneosa columellata, recorded for the second time by Esqueda et al.
(1998), 60 years after the holotype was registered from Arizona, USA (Long, 1941);
and Calvatia pygmaea, reported by Fries (1909) from Argentina and Bolivia, and found
again in Baja California by Ochoa et al. (1998).
MATERIALS AND METHODS
To analyze microscopical features of the basidiomata, freehand samples of the gleba
and peridium were mounted in water, lactophenol cotton blue, 5% KOH, and Melzer’s
reagent. Electronic micrographs were made under a Zeiss DSM 950 SEM following
Moreno et al. (1995). The material studied is deposited in the National Herbarium of the
Institute of Biology, UNAM, Mexico (MEXU), the mushroom collection of the Centro
de Estudios Superiores del Estado de Sonora, Mexico (CESUES), and the Herbarium
of the University of Alcalá, Madrid, Spain (AH).
1) Dpto. Biología Vegetal, Universidad de Alcalá, E-28871 Alcalá de Henares, Madrid, Spain; e-mail:
gabriel.moreno@uah.es; alberto.altes@uah.es
2) Centro de Investigación en Alimentación y Desarrollo, A. C. Apartado Postal 1735, Hermosillo,
Sonora, México, 83000; e-mail: esqueda@cascabel.ciad.mx
3) Laboratorio de Micología, Instituto de Biología, UNAM, Apartado Postal 70-233, Coyoacán,
México D. F., 04510; e-mail: psilva@ibiologia.unam.mx; therrera@ibiologia.unam.mx
© 2007 Nationaal Herbarium Nederland, Leiden University branch
P E R S O O N I A
Volume 19 / 2: 265–280 (Published 8 August 2007)

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SPECIES STUDIED
1. Agaricus texensis (Berk. & Curtis) Geml, Geiser & Royse — Figs. 1, 2
Agaricus texensis (Berk. & Curtis) Geml, Geiser & Royse, Mycol. Progr. 3 (2004) 172.
Secotium texense Berk. & Curtis in Berk., Grevillea 2 (1873) 34 – 35.
Gyrophragmium texense (Berk. & Curtis) Massee, Grevillea 19 (1891) 96.
Longia texensis (Berk. & Curtis) Zeller, Mycologia 35 (1943) 414.
Longula texensis (Berk. & Curtis) Zeller, Mycologia 37 (1945) 636.
The agaricoid basidiocarps have an obese and striate stalk, which extends as a per-
current columella through the pileus. The subglobose pileus remains closed until ma-
turity, when it expands to a broad, convex pileus similar to those of Agaricaceae. At
expansion, the peridium exposes the hymenophore, with its dark, crowded, wavy, free,
sometimes anastomosing lamellae; remains of the peridium frequently form a typical
double membranous annulus. Spores (statismospores) are subglobose to ovoid, smooth,
very dark, lacking a germ pore. Excellent illustrations of A. texensis are included in
Lloyd (1904c), Barnett (1943), Harding (1957), and States (1990).
The genus Longia was proposed by Zeller (1943), based on S. texense Berk. & Curtis
It was later abandoned on nomenclatural grounds, with the species transferred to the
new genus Longula (Zeller, 1945). It is closely related to G. dunalii (Fr.) Zeller, and
some authors (i. e. Guzmán & Herrera, 1969) considered both taxa as conspecific. Zeller
(1943) based their separation into two genera on account of the sturdiness of Longula
(Longia) and veil remnants at the base of the stipe of Gyrophragmium, features which he
compared with those of Amanita and Lepiota. Massee (1891) proposed the combination
G. texense, based on the presence of a volva in that species. Harding (1957) supports
that argument in his work on L. texensis var. major Zeller.
A recent molecular study has confirmed the differences between these two taxa (Geml
et al., 2004) based on sequences of the internal transcribed spacers (ITS) and partial
large subunit of ribosomal DNA, demonstrating that secotioid species G. dunalii and
L. texensis evolved from different species of Agaricus, and proposing their classification
in this genus as A. aridicola Geml, Geiser & Royse and A. texensis, respectively. How-
ever, the relationship between L. texensis and the genus Agaricus is not new: it has been
Figs. 1, 2. Agaricus texensis from Sonora (AH 31729). Spores under SEM. Scale bars: 1 µm.

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put forth by several authors and it is broadly accepted. In this sense, Barnett (1943) found
the development of L. texensis to be similar to that of some species of Agaricus.
Agaricus texensis (= L. texensis) is only known from the area between southwestern
United States and northwestern Mexico. It has been cited in Texas by Berkeley (1873)
as S. texense and by Lloyd (1904c) as G. texense; in New Mexico by Barnett (1943); in
California by Lloyd (1904b) as G. decipiens (Peck) Lloyd; in Oregon by Zeller (1943);
in Baja California by Ochoa et al. (1990) as G. dunalii (later corrected as L. texensis by
Ochoa, 1993). We have not examined the material studied by Urista et al. (1985), which
was recorded as G. dunalii; their description fits A. texensis instead. If this proposal is
accepted, its distribution would thus extend to the Mexican states of Nuevo León and
Coahuila. The only report of this species in Sonora was made by Ochoa (1993).
Specimens examined. Mexico: Sonora, La Colorada municipality, Km 31 Hermosillo to Yécora
road, microphyllous desert scrub, leg. M. Esqueda, A. Armenta, A. Núñez & R. Santos, 24.XI.1995,
CESUES 2215; ibidem, 16.II.1996, CESUES 2354; ibidem, 11.IX.1996, CESUES 2785. Km 40
Hermosillo to Yécora road, microphyllous desert scrub, leg. M. Esqueda, A. Armenta, A. Núñez &
R. Santos, 24.XI.1995, CESUES 2220; ibidem, 11.IX.1996, CESUES 2886. Km 100 Hermosillo to
Yécora road, tropical thorn forest, leg. M. Esqueda, A. Armenta, A. Núñez & R. Santos, 24.XI.1995,
CESUES 2230, duplo in AH 31728; ibidem, 14.V.1998, CESUES 3973; ibidem, 26.VIII.1998, CE-
SUES 4369, duplo in AH 31729.
2. Araneosa columellata Long — Figs. 3 –10
Araneosa columellata Long, Mycologia 33 (1941) 353.
Araneosa columellata produces closed gastroid carpophores, cream-coloured to
pale ochraceous, often pyriform, with a short and somewhat bulbous stipe as properly
illustrated by Long (1941). As he noted, the stipes of A. columellata detach easily from
the spore sac; the conic basal hole left by the stipe facilitates the breaking of the fragile
peridium, thus exposing the gleba. The Sonoran collections lack a stipe and have a
highly fragmented pileus. This is also found in the type collection of the species. As the
specific epithet reflects, a conspicuous columella protrudes through the pileus. The gleba
is formed by small, easily detachable peridioles, dark grey in the herbarium specimens,
surrounding the columella. The subglobose to irregularly ovoid spores are 5 – 6 (–7) ×
4 – 5(– 6) µm in diameter, pedicellate, reddish brown, smooth both under LM and SEM,
and without capillitium.
When Long (1941) created the genus Araneosa, he stated that A. columellata was
closely related to Arachnion Schwein. as the gleba is formed by small peridioles in both
cases. The latter does not possess a stipe nor columella. These two genera formed the
Arachniaceae, after being emended by Long himself (1941), who established a relation-
ship with Lycoperdaceae as in both families the gleba presents chambers until maturity.
With this similarity, Arachnion would be the closest genus to Lycoperdon (A. rufum
Lloyd and A. album Schwein.). Demoulin (1972) defended that relationship but did
not discuss the differences with Araneosa. To date, the Dictionary of Fungi (Kirk et
al., 2001) classifies Arachnion as Lycoperdaceae but leaves Araneosa in Agaricaceae.
The study and comparison of DNA sequences will allow to clarify the real affinities of
A. columellata.

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Figs. 3 –7. Holotype of Araneosa columellata (W.H. Long & V. Sandberg 7937 in BPI 736388).
3. Label of the collection; 4. basidiomes; 5 –7. spores under SEM. — Figs. 8 –10. Araneosa columellata
from Sonora (AH 31712). Spores under SEM. Scale bars: 4 = 10 mm; 5 –10 = 1 µm.
According to the only two reports of A. columellata (Long, 1941; Esqueda et al.,
1998) its distribution seems to be restricted to a small area between Nogales and Tuc-
son, Southern Arizona, and around Hermosillo, in central Sonora. This paper does not
change this range, but increases the scarce records of this species.

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Specimens examined. Mexico: Sonora, La Colorada municipality, Km 40 Hermosillo to Yécora
road, in Acacia constricta, Bursera microphylla and Guaiacum coulteri scrub, leg. M. Esqueda,
A. Armenta, A. Núñez & R. Santos, 14.X.1995, CESUES 2185, duplo in AH 31712. — USA: Arizona,
7 mi. from Nogales, in open mesquite flats, leg. W. H. Long & V. Sandberg 7937, 21.IX.1934, BPI
736388, holotype, and BR, isotype.
3. Calvatia bicolor (Lév.) Kreisel — Figs. 11, 12
Calvatia bicolor (Lév.) Kreisel, Persoonia 14 (1992) 435.
Bovista bicolor Lév., Ann. Sci. Nat., Bot. 5 (1846) 162.
Lanopila bicolor (Lév.) Pat., Bull. Soc. Mycol. France 15 (1899) 203.
Langermannia bicolor (Lév.) Demoulin & Dring, Bull. Jard. Bot. Belg. 45 (1975) 350.
Lasiosphaera fenzlii Reichardt in Fenzl, Reise Austr. Freg. Nov. 1 (1870) 135.
The Sonoran specimens are subglobose, approximately 50 × 70 – 80 mm. The dry exo-
peridium looks like a thin, fragile copper-coloured crust attached to the endoperidium,
which appears membranous and flexible, and is cream to pale ochre. Noticeable colour
variations of the exoperidium in some specimens are the origin of the specific epithet,
as pointed out by Bottomley (1948). The lack of subgleba and mycelial strands, and the
presence of the brown, cotton like, compact and persistent gleba (referring to the name
Lanopila), characterizes this species. The capillitium is abundant, dark, extensively
interwoven, c. 4 µm in diameter, somewhat branched, septate, not pitted. The spores
are yellow brownish, 5 – 6 µm in diameter, strongly ornamented, with SEM showing
almost cylindrical processes with rounded to almost planar tips, which sometimes form
small groups.
Calvatia bicolor may be clearly told apart from a Bovista when its gleba and the
features of the peridium are observed under the microscope, regardless of their macro-
scopic similarities. In any case, several authors have recorded this taxon under the genera
Bovista, Langermannia, Lanopila, or Lasiosphaera. This study follows Kreisel (1992),
who placed this taxon in genus Calvatia section Lanopila, but declines the generally
accepted conspecificity of C. bicolor and Lanopila wahlbergii Fr. (Homrich & Wright,
1973; Kreisel, 1994) until further study of the latter is made. The only sample of L. wahl-
bergii studied by us (Natal, South Africa, IX.1923, PREM 46086) has spores with spines
larger than those of C. bicolor, as observed with SEM.
Figs. 11, 12. Calvatia bicolor from Sonora (AH 31714). Spores under SEM. Scale bars: 1 µm.

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Calvatia bicolor is known from Argentina, Brazil, Ecuador, Cuba, St. Kitts (West
Indies), India, Indonesia (Lloyd, 1904a; Kreisel, 1994). It has also been cited from South
Africa and the former Belgian Congo by Bottomley (1948). We maintain our doubts
about some identifications cited as L. wahlbergii until further information is acquired.
The first reference (and illustration) of this species in Mexico was made by Lloyd
(1902) as B. lateritia Berk., and corrected later by the author (Lloyd, 1904a). Guzmán
& Herrera (1969) cited L. wahlbergii from the Mexican state of Morelos. Ochoa (1993)
included two collections of L. bicolor from Baja California Sur in his doctoral thesis
(unpublished). The material studied for the present work is the first record of C. bicolor
for Sonoran mycobiota.
Specimens examined. Mexico: Sonora, Baviácora municipality, near Baviácora town, leg. M. Es-
queda & R. Ramírez, 26.III.1993, CESUES 1220, duplo in AH 31714; Ónavas municipality, Km 204.5
Hermosillo to Yécora road, oak-pine forest, leg. M. Esqueda, A. Armenta, A. Núñez & R. Santos,
6.X.1995, CESUES 2108, duplo in MEXU 24540; ibidem, leg. E. Pérez-Silva, T. Herrera, M. Es-
queda, A. Armenta, A. Núñez & R. Santos, 13.IX.1996, CESUES 2973; San Javier municipality, Km
151 Hermosillo to Yécora road, tropical thorn forest, leg. M. Esqueda, A. Armenta, A. Núñez & R. San-
tos, 14.X.1995, CESUES 2147.
4. Calvatia craniiformis (Schwein.) Fr. ex De Toni — Figs. 13, 14
Calvatia craniiformis (Schwein.) Fr. ex De Toni in Saccardo, Syll. Fung. 7 (1888) 106.
Bovista craniiformis Schwein., Trans. Amer. Philos. Soc., ser. 2, 4 (1832) 256.
The type species of Calvatia is characterized by a cellular and well-developed sub-
gleba, resembling C. cyathiformis (Bosc) Morgan and H. utriformis (Bull.: Pers.) Kreisel.
Calvatia craniiformis differs from the former by the olivaceous, not lilacineous gleba,
and by the smaller spores (3 – 4 µm in diameter), semi smooth with LM but with small
and regularly distributed verrucae-spines as observed with SEM. The capillitium is
conspicuously pitted, with holes up to 3 µm in diameter, whereas the European species
H. utriformis has the characteristic capillitium of genus Handkea (Kreisel, 1989) with
elongated, slit-like pits. In addition, even when the spores of the latter are semi smooth
under LM, the ornamentation under SEM appears more dense with rounded verrucae.
Figs. 13, 14. Calvatia craniiformis from Sonora (AH 31715). Spores under SEM. Scale bars: 1 µm.

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The distribution of C. craniiformis includes North America and areas as remote as
China and Japan (Kreisel, 1994). This species is well known in the United States, as
proven by the numerous specimens kept in the Herbarium NY. However, the records
from that country are rather scarce: North Carolina (type), Michigan (Kreisel, 1994),
Oklahoma, Texas (Zeller & Smith, 1964). On the other hand, it is not a common species
in Mexico, cited only from the northern states of Nuevo León, Coahuila (Urista et al.,
1985) and Sonora (the two collections studied here).
Recently, Coetzee & Van Wyk (2003) wrote a note explaining the problem related
to the authory of the name C. craniiformis.
Specimens examined. Mexico: Sonora, Yécora municipality, Km 251 Hermosillo to Yécora road,
oak forest, leg. M. Esqueda & R. Rodríguez, 10.IX.1995, CESUES 1922; ibidem, leg. E. Pérez-Silva,
T. Herrera, M. Esqueda, A. Armenta & R. Santos, 13.IX.1996, CESUES 2977, duplo in AH 31715.
5. Calvatia pygmaea (R. E. Fr.) Kreisel, G. Moreno, C. Ochoa & Altés — Figs. 15,
16
Calvatia pygmaea (R. E. Fr.) Kreisel, G. Moreno, C. Ochoa & Altés, Cryptog. Mycol. 19 (1998)
132.
Lanopila pygmaea R. E. Fr., Ark. Bot. 8 (1909) 16 –17.
Calvatia pygmaea is an interesting xerophilous bovistoid species, which was practi-
cally unnoticed until recently. Very few records are available: the two collections from
Argentina and Bolivia which allowed Fries (1909) to describe it, and another two sam-
ples collected by Ochoa et al. (1998) in Baja California Sur, Mexico. The first record
from Sonora of C. pygmaea is hereby presented, where it appears to be a relatively
frequent species in sandy basic soils under Bursera, Cercidium, and Olneya.
Calvatia pygmaea belongs to Calvatia section Lanopila (Fr.) Kreisel, but it is the only
species in the section showing smooth spores with both LM and SEM. The collections
from Sonora have globose to subglobose basidiocarps with a diameter of 10 – 25 mm
and without subgleba. Its features, including the size of spores 6.5 – 8 µm in diameter,
fit very well the original description and the above-mentioned collections (Ochoa et al.,
1998).
Figs. 15, 16. Calvatia pygmaea from Sonora (AH 31717). Spores under SEM. Scale bars: 1 µm.

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Specimens examined. Mexico: Sonora, Hermosillo municipality, Km 115 36N St. to Puerto Lib-
ertad road, microphyllous desert scrub, leg. M. Esqueda, A. Armenta, M. Coronado & A. Sánchez,
7.III.1998, CESUES 3953, duplo in AH 31717; ibidem, 29.VIII.1998, CESUES 4373, duplo in AH
31718; Km 125 36N St. to Puerto Libertad road, microphyllous desert scrub, leg. M. Esqueda, A.
Armenta, M. Coronado & A. Sánchez, 29.VIII.1998, CESUES 4401, duplo in AH 31716; Ejido
6 de Diciembre, leg. G. Yanez, 27.VII.1994, CESUES 3257, duplo in AH 31719; Puerto Peñasco
municipality, Pinacate and Grand Desert Biosphere Reserve, Ejido Los Norteños, leg. A. Sánchez,
I. Encinas & J. Miranda, 12.VIII.2004, CESUES 5248; Sonoyta municipality, Pinacate and Grand
Desert Biosphere Reserve, Ejido Punta Peñasco, leg. A. Sánchez, I. Encinas & J. Miranda, 16.I.2004,
CESUES 5102; ibidem, 28.IV.2004, CESUES 5187; ibidem, 12.VIII.2004, CESUES 5213.
6. Disciseda hyalothrix (Cooke & Massee) Hollós — Figs. 17, 18
Disciseda hyalothrix (Cooke & Massee) Hollós, Növényt. Közlem. 1 (1902) 107.
Bovista hyalothrix Cooke & Massee, Grevillea 16 (1888) 73.
Catastoma hyalothrix (Cooke & Massee) Lloyd, The Lycoperdaceae of Australia, New Zealand
and Neighbouring Islands (1905) 27.
Catastoma pedicellatum Morgan, J. Cincinnati Soc. Nat. Hist. 14 (1892) 143 –144.
Disciseda pedicellata (Morgan) Hollós, Természetrajzi Füz. 25 (1902) 103.
The Sonoran specimens of D. hyalothrix have basidiocarps up to 20 mm in diameter.
This species is characterized by ochraceous spores of a variable size (8.5 –13 µm in
diameter), with conspicuous ornamentation formed by spines fused at the apex, forming
processes with a flat tip. The spores maintain pedicels not as often as in the collections
studied by Moreno et al. (2003), with a length of 5 µm, which in some cases reaches up
to 12 µm. We believe that it is the grade of maturity and conservation of the gleba that
determines such a condition, and that the spore ornamentation is much more important
for the characterization of the species.
Recently, a comparative study of the type collections of D. hyalothrix and D. pedi-
cellata (Moreno et al., 2003) showed their conspecificity. Unfortunately, the name to
be given priority is the least known: the name D. hyalothrix has been applied only to
specimens from some Australian localities. Previously, Grgurinovic (1997) established
the synonymy of B. irregularis Berk. and D. pedicellata and proposed a new species at
the same time, D. errurraga, which Grgurinovic identified earlier as D. hyalothrix, based
Figs. 17, 18. Disciseda hyalothrix from Sonora (AH 31722). Spores under SEM. Scale bars: 1 µm.

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Moreno, Esqueda, Pérez-Silva, Herrera & Altés: Fungi from Sonora, Mexico
on Australian collections. In both cases, the descriptions emphasize some differences
in spore size and pedicel length, just the characters that are now considered to be the
most variable. To date, we have had no access to the type collections of B. irregularis
or D. errurraga, but we believe that study of their spore ornamentation could lead to
the conclusion that these names are additional synonyms of D. hyalothrix. In case this is
confirmed, the name D. hyalothrix would be correct because its basionym, B. hyalothrix,
was published earlier (March 1888) than B. irregularis (May 1888).
This species has been collected in almost all the continents, usually under the name
of D. pedicellata. Even though the number of published records is not very significant,
we cite here: USA (Coker & Couch, 1928), Argentina (Spegazzini, 1912; Domínguez de
Toledo, 1989), Australia (Cunningham, 1942; Grgurinovic, 1997), Mozambique, South
Africa (Bottomley, 1948), and a very doubtful citation from Sweden (Rydberg, 1949;
Eckblad, 1955). Some records of this species are from xeric areas in several states of
northern Mexico: Chihuahua (Laferrière & Gilbertson, 1992), Sonora (Esqueda et al.,
1995), and Baja California (Ochoa & Moreno, 1996; Ochoa et al., 2000).
Specimens examined. Mexico: Sonora, Hermosillo municipality, Centro Ecológico de Sonora, leg.
M. Esqueda & R. E. Villegas, 7.V.1993, CESUES 1581, duplo in AH 31721; Km 125 Hermosillo to
Puerto Libertad road, leg. M. Esqueda, A. Armenta, A. Núñez & A. Sánchez, 29.VIII.1998, CESUES
4402, duplo in AH 31720; La Colorada municipality, Km 100 Hermosillo to Yécora road, leg.
E. Pérez-Silva, M. Esqueda, A. Armenta, A. Nuñez & R. Santos, 11.IX.1996, CESUES 2842; San Javier
municipality, Km 137.5 Hermosillo to Yécora road, tropical deciduous forest, leg. M. Esqueda,
A. Armenta, A. Núñez & R. Santos, 9.VI.1996, CESUES 2768, duplo in AH 31722.
7. Disciseda stuckertii (Speg.) G. Moreno, Esqueda, Altés, comb. nov. — Figs. 19 –
21
Bovista stuckertii (‘stuckerti’) Speg., Anales Mus. Nac. Buenos Aires 8 (1902) 56.
Abstoma stuckertii (Speg.) J. E. Wright & V. L. Suárez, Cryptog. Bot. 1 (1990) 377.
The genus Abstoma was proposed by Cunningham (1926) for Lycoperdaceae resem-
bling Bovista or Disciseda, basidiocarps with an irregular dehiscence, without a definite
stoma, wavy capillitium without pits and reticulate spores. Later, it was emended by
Wright & Suárez (1990), including smooth and ornamented spores. Finally, it was
corrected by Moreno et al. (1992) to include the new species A. friabile (‘friabilis’)
G. Moreno et al. with a distinctive pitted capillitium. According to Wright & Suárez
(1990) there are four species with reticulate spores: A. purpureum (Lloyd) G. Cunn.
(New Zealand), A. townei (Lloyd) Zeller (West and Southwest United States, West
Argentina), A. stuckertii (Argentina) and A. reticulatum G. Cunn. (Australia, West and
Southwest United States). Among these species, the size and ornamentation of spores are
so similar that clear limits are difficult to set. On this basis, Domínguez de Toledo (1989)
established in her doctoral thesis the conspecificity of A. stuckertii and A. reticulatum.
The Mexican collections studied met all the characteristics in this group of Abs-
toma species, especially those from the center of the country which were identified as
D. muelleri (Berk.) G. Cunn. by Guzmán & Herrera (1969). Spores in the collections
from Sonora are slightly smaller (7–10 µm in diameter) and meshes per hemisphere are
less numerous, but do not exceed the ranges observed in such species. Pending a solution
to the possible synonymy of these species with reticulate spores, our initial plan was to
classify the Mexican material as A. stuckertii, the name with priority.

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On the other hand, the suitability of maintaining the genus Abstoma should be recon-
sidered given the similarities with other close genera, especially Disciseda (absence of
subgleba, sinuous-spiraled capillitium, similar size and spore morphology). The main
difference between these two genera is reduced to the type of dehiscence, by a more or
less defined stoma in Disciseda, and by an apical irregular fracture of the spore sac in
Abstoma. But this character is very variable in the collections of Abstoma studied by
us, probably determined by the age and conservation of the material. Molecular data
will most likely provide accurate information leading to a more adequate placement
of some species of Abstoma in the genus Disciseda, as we do here, and perhaps some
other species (i. e. A. friabile) can better be placed in Calvatia.
Regarding A. stuckertii, in both the Mexican material and the Argentinian collection
BAFC 31871, it was proven that dehiscence is produced by a more or less defined stoma.
Given the sand case of exoperidium, added to the most distinctive characteristic of
Disciseda, we deem reasonable to propose a new combination: D. stuckertii. However,
the link between A. stuckertii and Disciseda is not new at all. The species was origi-
nally classified by Spegazzini (1902) as B. stuckerti, but years later (Spegazzini, 1912)
he reconsidered that position and placed that name in the synonymy of D. pampeana
(Speg.) Speg. (= B. pampeana Speg.). Now we know that such synonymy is inaccu-
rate because of the obvious differences in spore ornamentation of the two species (cf.
Wright & Suárez, 1990), but the sand case that we mentioned before seems to us to be
an outstanding character. Later, Wright & Suárez (1990) proposed the new combina-
tion of this species as A. stuckertii after revising genus Abstoma in South America, and
accepted the synonymy of D. pila R.E. Fr., which was suggested earlier by Spegazzini
himself (1912).
Wright & Suárez (1990) reported Disciseda stuckertii from several provinces of
Argentina: Chaco, Córdoba (type of Bovista stuckertii), Jujuy (type of Disciseda pila),
and Santiago del Estero. These authors also included the only non-Argentinian record,
that is from New Zealand. Therefore, the material studied here represents the first record
of D. stuckertii for Mexico.
Specimens examined. Mexico: Sonora, Hermosillo municipality, Centro Ecológico de Sonora,
microphyllous desert scrub, R. E. Villegas & M. Esqueda, 7.V.1993, CESUES 1290; ibidem,
Figs. 19 – 21. Disciseda stuckertii from Sonora (AH 31709). Spores under SEM. Scale bars: 1 µm.

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Moreno, Esqueda, Pérez-Silva, Herrera & Altés: Fungi from Sonora, Mexico
20.VII.1993, CESUES 1587; La Colorada municipality, Km 100 Hermosillo to Yécora road, open
areas in tropical thorn forest, leg. A. Armenta, A. Núñez & R. Santos, 16.II.1996, CESUES 2363, duplo
in AH 31709; Baviácora municipality, Km 72 Mazocahui to Hermosillo road, leg. E. Pérez-Silva,
T. Herrera & M. Esqueda, 30.VIII.2001, CESUES 4842; Sonoyta municipality, Pinacate and Grand
Desert Biosphere Reserve, Papalote, leg. A. Sánchez, I. Encinas & J. Miranda, 12.VIII.2004, CESUES
5232; Distrito Federal, Tranvia way, Xochimilco to Manantiales, leg. G. Guzmán 293, 13.IX.1955,
ENCB, duplo in AH 31710; Estado de México, Zumpango, edge of big drainage channel, leg.
G. Guzmán 316, 18.IX.1955, ENCB, duplo in AH 31711.
8. Disciseda verrucosa G. Cunn. — Figs. 22, 23
Disciseda verrucosa G. Cunn., Trans. & Proc. New Zealand Inst. 57 (1926) 205.
Disciseda arida Velen., Novit. Mycol. (1939) 169.
Disciseda verrucosa is clearly characterized by its sparsely pedicellate spores con-
spicuously ornamented by obtuse finger-like processes, usually curved at the apex.
Recently, Moreno et al. (2003) studied the type material of D. verrucosa and D. arida,
and proposed them as conspecific. Even considering this previous study, this taxon
has been scarcely cited up to present; some records are known from New Zealand
and Australia (Cunningham, 1942), South Africa (Bottomley, 1948), from the former
Czechoslovakia (Moravec, 1958), and Spain (Moreno et al., 2003). Sonora is the only
site in Mexico from which D. verrucosa is reported. The material studied adds to the
previous records by Aparicio-Navarro et al. (1994) and Pérez-Silva et al. (2000).
Specimens examined. Mexico: Sonora, Hermosillo municipality, Las Dunas, close to Puerto
Libertad town, leg. A. Armenta, A. Núñez & A. Sánchez, 29.VIII.1998, CESUES 4484; La Colorada
municipality, Km 100 Hermosillo to Yécora road, leg. M. Esqueda, A. Armenta, A. Núñez & R. Santos,
17.III.1996, CESUES 2615a; ibidem, 2.VII.1997, CESUES 3299, duplo in AH 31648; San Javier
municipality, Km 151 Hermosillo to Yécora road, leg. A. Armenta, A. Núñez & R. Santos, 31.V.1996,
CESUES 2711; Yécora municipality, 3.4 km Yécora to Las Cabañas way, leg. M. Esqueda &
R. Rodríguez, 8.IX.1995, CESUES 1778; Sonoyta municipality, Pinacate and Grand Desert Biosphere
Reserve, Papalote, leg. A. Sánchez, I. Encinas & J. Miranda, 12.VIII.2004, CESUES 5226.
Figs. 22, 23. Disciseda verrucosa from Sonora (AH 31648). Spores under SEM. Scale bars: 1 µm.

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Moreno, Esqueda, Pérez-Silva, Herrera & Altés: Fungi from Sonora, Mexico
9. Endoptychum arizonicum (Shear & Griffiths) Singer & A. H. Sm. — Figs. 24 –
32
Endoptychum arizonicum (Shear & Griffiths) Singer & A. H. Sm., Brittonia 10 (1958) 221.
Secotium arizonicum Shear & Griffiths in Shear, Bull. Torrey Bot. Club 29 (1902) 450 – 451.
This taxon is frequently observed in the xerophilous areas of Sonora. It produces
solitary to scattered, small basidiocarps (10 –13 mm in diameter in our collections),
resembling an immature Bovista. Endoptychum arizonicum is characterized by a
membranous, rather thin, smooth, yellowish exoperidium, fragile when dry; the stipe
is very short or practically absent, continued by a not fully developed columella; the
gleba is sublamellar, consisting of closely anastomosed, thin, yellowish tramal plates;
the spores are smooth, thick walled, subhyaline, globose, 7–12 µm in diameter, with a
short pedicel, and germ pore absent. The presence of clamp-connections in this species
(specially in the columella), pointed out only by Vellinga et al. (2003) was confirmed.
Except for the rather smaller basidiocarps, the characteristics of the Mexican material
match those we observed in various collections from Tucson (Arizona, USA), including
the type and isotype specimens, Griffiths 1630 and West American Fungi 323, respec-
tively (Shear, 1902). Amongst the materials from Arizona that we have studied, there
are two other samples (Lloyd 30824 and Griffiths 205) mislabelled as type.
Endoptychum arizonicum was first considered to be very close to E. agaricoides
Czern. (Shear, 1902; Lloyd, 1903). There are actually many differences between the
two species: E. agaricoides has larger, scaly basidiocarps; a percurrent, well-developed
columella; ellipsoidal spores not larger than 10 µm in diameter, greenish to yellowish
brown, with germ pore. Some other species, with obvious differences such as darker
spores, have been erroneously included in this genus, i. e. E. depressum Singer & A.H.
Sm. from North America, and E. melanosporum (Berk.) Singer & A. H. Sm. (= S. melano-
sporum Berk.) from Australia. In 1997, Grgurinovic proposed three new species from the
Australian material: E. wariatodes Grgur., E. kolya Grgur., and E. moongum Grgur.).
The molecular data of Endoptychum lead us to believe that some of the species
recorded under this genus should be classified elsewhere. Vellinga et al. (2003) demon-
strated a close relationship between E. agaricoides, C. molybdites (G. Mey.: Fr.) Massee,
and some species of Macrolepiota; Singer & Smith (1958) had suggested this based on
the greenish colour of their spores. Vellinga et al. (2003) proposed to group these taxa
in the genus Chlorophyllum. That would entail conserving the name Chlorophyllum
over Endoptychum (Vellinga & de Kok, 2002), in spite of the latter being much older;
a solution from the Committee for Fungi is still pending. During the process of reviewing
and publishing of this paper, the Committee for Fungi has accepted the proposal by Vellinga
& de Kok (2002) and recommends to conserve Chlorophyllum over Endoptychum
H
Figs. 24, 25. Endoptychum arizonicum from Arizona (D. Griffiths 205 in BPI 728601). 24. Label of
the collection; 25. basidiomes. — Fig. 26. Holotype of Endoptychum arizonicum (D. Griffiths 1630
in BPI 797001). Spore under SEM. — Figs. 27, 28. Isotype of Endoptychum arizonicum (D. Griffiths
323 in BPI 728540). Spores under SEM. — Figs. 29 – 32. Endoptychum arizonicum from Sonora.
29. Sectioned basidiome (AH 31725); 30. basidiomes (AH 31723); 31, 32. spores under SEM (AH
31725). Scale bars: 25, 29, 30 = 10 mm; 26 – 28, 31, 32 = 1 µm.

PERSOONIA — Vol. 19, Part 2, 2007
278
(Gams, 2005; Taxon 54: 520–522). This would result in combinations like C. agari-
coides (Czern.) Vellinga (Vellinga, 2002). Following the same line, E. depressum is
apparently related to Agaricus arvensis Schaeff., as Singer & Smith suggested (1958),
and would be included in genus Agaricus as A. inapertus Vellinga (Vellinga et al., 2003).
It will be interesting to include molecular data to solve the position of E. arizonicum
and the other species of the genus.
Endoptychum arizonicum was found to be a characteristic species of the xerophi-
lous areas between the United States and Mexico (Shear, 1902; Arora, 1986; Guzmán
& Herrera, 1969; Esqueda et al., 1998). However, its distribution does not appear to
be restricted to that region, considering the records from Argentina (Singer, 1962) and
more recently from Italy (Sarasini & Contu, 2001).
Specimens examined. Mexico: San Luis Potosí, Tanque Caballos, Km 34 San Luis Potosí to
Antiguo Morelos road, leg. F. Medellín-Leal, 20.VI.1958, ENCB, duplo in AH 31726; Sonora, Altar
municipality, Monte La Esperanza, 4 km NE Altar town, leg. H. Araiza, 18.IX.1968, ENCB, duplo
in AH 31727; La Colorada municipality, Km 31 Hermosillo to Yécora road, microphyllous desert
scrub, leg. M. Esqueda, A. Armenta, A. Núñez, R. Rodríguez & R. Santos, 14.X.1995, CESUES
2204, duplo in AH 31723; ibidem, 24.XI.1995, CESUES 2210; ibidem, 16.II.1996, CESUES 2351;
ibidem, 29.IV.1996, CESUES 2707; ibidem, 11.IX.1996, CESUES 2789, duplo in AH 31724; Km 40
Hermosillo to Yécora road, microphyllous desert scrub, leg. M. Esqueda, A. Armenta, A. Núñez &
R. Santos, 14.X.1995, CESUES 2176; ibidem, 24.XI.1995, CESUES 2221; ibidem, 29.IV.1996,
CESUES 2705; ibidem, 11.IX.1996, CESUES 2843, duplo in AH 31725; ibidem, leg. M. Esqueda,
2.VII.1997, CESUES 3305; Km 100 Hermosillo to Yécora road, tropical thorn forest, leg. M. Es-
queda, A. Armenta, A. Núñez & R. Santos, 29.IV.1996, CESUES 2696; Puerto Peñasco municipality,
Pinacate and Grand Desert Biosphere Reserve, Ejido Los Norteños, leg. M. Esqueda, M. Coronado,
A. Sánchez, J. Miranda & I. Encinas, 28.IV.2004, CESUES 5189; Sonoyta municipality, Pinacate
and Grand Desert Biosphere Reserve, Ejido Punta Peñasco, leg. M. Esqueda, A. Sánchez, I. Encinas
& J. Miranda, 28.IV.2004, CESUES 5193. — USA: ArizonA, leg. D. Griffiths 205, X.1905, BPI
728601, ‘type’ in label (?); Tucson, leg. D. Griffiths, Herb. Lloyd 30824, BPI 718209, ‘type’ in label
(?); ibidem, leg. D. Griffiths, IX.1930, Herb. Long 7895, BPI 728538; ibidem, leg. D. Griffiths 1630,
IX.1900, BPI 797001, holotype; ibidem, leg. D. Griffiths 323, IX.1900, BPI 728540 & 796870, iso-
types; Nogales-Tucson road, 7 miles from Tucson, 19.II.1934, Herb. Long 7629 (BR); Catlaw Flats,
4 miles from Sabino Canyon, near Tucson, 28.IX.1939, Herb. Long 8403 (BR); unknown locality,
Herb. Long 7854 (BPI 728539).
ACKNOWLEDGEMENTS
The authors thank SEMARNAT-CONACYT (Grant 2002-C01-0409) (México); DGAPA IN206901
UNAM (México); ‘Ministerio de Ciencia y Tecnología, Plan Nacional de Investigación Científica,
Desarrollo e Innovación Tecnológica REN2002-01965’ and ‘Agencia Española de Cooperación In-
ternacional’ (AECI) (Spain) for financial support. Our gratitude goes to Dr. F. Chiang (UNAM) and
Prof. R. P. Korf (Cornell University) for improving the English version, and the latter one, furthermore,
for his nomenclatural suggestions. Thanks to J. A. Pérez and A. Priego, of the SEM Service of the
University of Alcalá, for their assistance.
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