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ISSN (print) 0093-4666 © 2015. Mycotaxon, Ltd. ISSN (online) 2154-8889
MYCOTAXON
http://dx.doi.org/./.
Volume 130, pp. 145–164 January–March 2015
Myxomycetes of Sonora (Mexico) 6.
Central plains of the Sonoran Desert
M L , G M *, M E ,
C S-M , M L. C
1 Dpto. Ciencias Químico Biológicas, Instituto de Ciencias Biomédicas, Univ. Autónoma de Ciudad Juárez,
Anillo Envolvente Pronaf y Estocolmo s/n, Ciudad Juárez, Chihuahua 32300, México
2 Dpto. Ciencias de la Vida, Edicio de Biología, Univ. Alcalá, Madrid, 28805, España
3 Centro de Investigación en Alimentación y Desarrollo,
A.C. Apartado Postal 1735, Hermosillo, Sonora, 83304, México
4 Univ. Estatal de Sonora. Apartado Postal 11, Admón. 11, Hermosillo 83000, Sonora, México
* C : gabriel.moreno@uah.es
A — Myxomycetes from 10 localities in the central plains of the Sonoran desert were
studied using the moist chamber culture technique. irty-three species have been identied,
of which ve (Comatricha mirabilis, Cribraria confusa, Didymium eremophilum, Perichaena
luteola, and P. stipitata) represent new records for Sonora, and Comatricha mirabilis is cited
for the rst time from Mexico. Light and scanning electron microphotographs illustrate the
most diagnostic characters of these species, which are rare or scarcely mentioned in the
literature.
K — Amoebozoa, chorology, myxobiota, slime moulds, taxonomy
Introduction
e Sonoran desert, which lies in northwestern Mexico and the southwestern
United States, covers a 260,000-km2 area, with the largest portion occurring in
Sonora state (Mexico) and the rest in southwestern Arizona and southeastern
California (United States). is is the warmest and the most arid North
American desert due to its low elevation (≤600 m) and 22–24 °C average annual
temperature. e bi-seasonal (winter and summer) rainfall averages 200–400
mm per year (Hernández 2006).
e desert vegetation is characterized by trees and shrubs, including
Parkinsonia orida (A. Gray) S. Watson, Olneya tesota A. Gray, Prosopis
glandulosa Torr., Fouquieria macdougalii Nash, and Bursera spp. e majority
of the cacti are large, with Carnegiea gigantea (Engelm.) Britton & Rose the
146 ... Lizárraga & al.
T 1. Studied Sonoran localities and types of vegetation
L L N L W A V
Town of Benjamin Hill
1. Las Animas 30°12′37.95″111°18′55.04″770 m MDS
Town of Opodepe
2. Tuape 30°02′52.89″111°00′22.05″670 m M
Town of Guaymas
3. Cañón de Nacapule 27°59′08.54″111°02′06.40″81 m SC
4. El Tigre 28°06′31.14″111°01′21.11″139 m SC
5. El Apache 28°19′25.33″111°14′27.74″43 m M
6. Maytorena 28°13′32.49″110°48′50.28″40 m SC
Town of Carbó
7. San Luis 29°33′42.38″111°04′52.79″458 m MDS
Town of Hermosillo
8. La Primavera 28°48′06.37″111°09′09.62″167 m MDS
9. La Pintada 28°39′56.35″110°57′15.64″233 m SC
10. El Papalote 29°12′56.14″111°02′39.43″348 m M
Types of vegetation: MDS = microphyllous desert scrub; SC = sarcocaule scrub; M = mezquital
most representative (Hernández 2006). Myxomycetes can develop in arid
environments due to decomposition of and water retained by succulent plants
such as Opuntia spp. ese provide an excellent microhabitat with enough
moisture for optimal development. Although the Sonoran desert has a high
level of biological diversity, here have been only a few studies of its myxobiota.
e Sonoran desert records of Evenson (1961) and Blackwell & Gilbertson
(1980a,b; 1984) were obtained mainly in Arizona (U.S.A.) in the United States.
Although there have been various diverse studies of the myxobiota in Sonora
(Mexico), only twenty-seven reports are known for the biosphere reserve
El Pinacate and Gran Desierto de Altar (Esqueda et al. 2013, Moreno et al.
2004); most studies have been centered in pine–oak forests and lowland areas
(Pérez-Silva et al. 2001; Moreno et al. 2006; Lizárraga et al. 2007, 2008a,b).
e catalogue of myxomycetes for Sonora state (Moreno et al. 2007) has
been increased from 77 species to 108 species by Lizárraga et al. (2007, 2008a,b)
and Esqueda et al. (2013). e present contribution brings the total number to
113.
Materials & methods
Four surveys were conducted in 2009, one each season of the year, in 10 localities
of the central plains of the Sonoran desert (T 1). Specimens were gathered or
cultured in moist chamber cultures. Permanent slides were prepared in Hoyer’s liquid
for microscopic study. Collections have been deposited in the collection of fungi of the
Myxomycetes of Sonora 6 (Mexico) ... 147
Universidad Estatal de Sonora, Hermosillo, Sonora, México (UES), with a small part in
the herbarium of the Universidad de Alcalá, Alcalá de Henares, Madrid, Spain (AH).
e spore ornamentation was examined under SEM using the material deposited in
AH. Collector names are abbreviated as follows: AG = A. Gutiérrez; ML = M. Lizárraga;
CS = C. Salazar; and AS = A. Sánchez. New species records for Sonora are marked with a
single asterisk, and the species recorded as new for Mexico is marked with two asterisks.
Taxonomy
Arcyria cinerea (Bull.) Pers.
S : MEXICO. S: Loc. 4, on Forchhammeria watsonii Rose,
leg. ML, CS, AS & AG, 19-VIII-2009, obtained from moist chamber, 30-IX-2009 (UES
8539); obtained from moist chamber, 9-X-2009 (UES 8541).
C— Previously cited for Sonora by Lizárraga et al. (2007, 2008b).
Arcyria denudata (L.) Wettst.
S : MEXICO. S: Loc. 1, on decomposing wood, leg. AG & AS,
22-V-2009, obtained from moist chamber, 11-VI-2009 (UES 8106).
C—Previously recorded for Sonora by Pérez-Silva et al. (2001), and
Lizárraga et al. (2008a,b).
Badhamia gracilis (T. Macbr.) T. Macbr.
S : MEXICO. S: Loc. 1, on Cylindropuntia sp., leg. AS & AG,
9-III-2009, obtained from moist chamber, 20-IV-2009 (UES 7858); Loc. 2, on Opuntia
sp., leg. AS & AG, 14-III-2009, obtained from moist chamber, 15-IV-2009 (UES 7854);
Loc. 3, on Stenocereus thurberi (Engelm.) Buxb., leg. AS & AG, 26-V-2009, obtained
from moist chamber, 30-VI-2009 (UES 8206); Loc. 4, on Lophocereus schottii (Engelm.)
Britton & Rose, leg. CS, AS & AG, 7-XI-2009, obtained from moist chamber, 4-XII-2009
(UES 8660); Loc. 5, on Parkinsonia sp., obtained from moist chamber, 20-IV-2009 (UES
7856); Loc. 6, on Cylindropuntia versicolor (Engelm.) F.M. Knuth, leg. AG & AS, 12-
III-2009, obtained from moist chamber, 6-IV-2009 (UES 7845); Loc. 7, on Stenocereus
thurberi, obtained from moist chamber, 14-IV-2009 (UES 7962); Loc. 8, on Lophocereus
schottii, leg. AS & AG, 14-III-2009, obtained from moist chamber, 14-IV-2009 (UES
7847); Loc. 9, on Cylindropuntia arbuscula (Engelm.) F.M. Knuth, leg. AG & AS 26-V-
2009, obtained from moist chamber, 11-VI-2009 (UES 8110); Loc. 10, on Parkinsonia
sp., leg. AS & AG, 12-III-2009, obtained from moist chamber, 20-IV-2009 (UES 7860).
C—We follow the taxonomy of Moreno & Oltra (2010), who regarded
Badhamia melanospora Speg. as separate from B. gracilis. e two species dier
mainly in spore morphology, with the spores >15–17 µm diam., polyhedral,
and densely warty in B. melanospora and <12–14 µm diam, polyhedral, and
with scattered warts with intervening spaces in B. gracilis. Both species fruit
mainly on succulent plants (Castillo et al. 1996). Some studied collections were
mistakenly identied as B. melanospora by Moreno et al. (2006) and Esqueda
et al. (2013).
148 ... Lizárraga & al.
Badhamia spinispora (Eliasson & N. Lundq.) H.W. Keller & Schokn. F , –
Fructications sessile, subglobose to short plasmodiocarps, white to
grey, growing grouped together. Sporotheca 0.2–1 × 0.2–0.5 mm. Peridium
double, calcareous, with irregular dehiscence. Hypothallus inconspicuous.
Pseudocolumella irregular, calcareous. Capillitium scarce and joined by
strands consisting of irregular calcareous nodules. Spores (9–)10–12 × 13–15
µm, globose to subglobose, black in mass, violaceous with a paler area under
LM, ornamented with large spines. SEM shows that the spore ornamentation is
formed by irregularly scattered or grouped <0.5 µm high warts.
S : MEXICO. S: Loc. 9, over hare manure, leg. AG & AS, 26-V-
2009; obtained from moist chamber, 30-VI-2009 (UES 8116, AH 42974).
C—is species is characterized by its generally coprophilous habitat
(although it does occasionally grow on tree bark), cespitose habit, capillitium
that varies from physaroid to badhamioid, and spores that are dark in mass,
spiny, and with a paler area. A study of B. spinispora, Physarum apiculosporum
Härk., and Badhamia verrucospora G. Moreno & al. was published by Moreno
et al. (2011).
Cited from Sonora by Esqueda et al. (2012, 2013).
Comatricha elegans (Racib.) G. Lister
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AG & AS, 22-V-
2009, obtained from moist chamber, 3-VI-2009 (UES 8120); Loc. 7, leg. ML, CS, AS &
AG, 17-VIII-2009, obtained from moist chamber, 7-IX-2009 (UES 8563); Loc. 9, 20-
VIII-2009, obtained from moist chamber, 9-IX-2009 (UES 8621); Loc. 10, decomposing
wood, 21-V-2009, obtained from moist chamber, 11-VI-2009 (UES 8122).
C—Lizárraga et al. (2005) published a study (including SEM
photomicrographs) of C. elegans from Chihuahua.
Previously recorded from Sonora by Lizárraga et al. (2007, 2008a).
**Comatricha mirabilis R.K. Benj. & Poitras F 7–12
Fructications stipitate, 0.7–0.9 mm tall. Sporotheca subglobose to
subcylindrical, 0.4–0.6 mm diam. Peridium eeting and persistent at the
base, resembling a necklace. Stipe short, dark, 0.3–0.6 mm high. Hypothallus
membranous, hyaline, as common in Comatricha. Columella crossing the
sporotheca and forking into several branches at the apex. Capillitium parallel
along the columella, dark brown with free ends, dichotomous, widened and
forked at the tips that protrude at the periphery of the sporotheca. Spores dark
brown in mass, 9–12 µm diam., globose to subglobose, reticulate, with walls
1–2 µm high. SEM observation reveals that the spore is reticulate and the high
banded reticulum has entire and unperforated walls.
Myxomycetes of Sonora 6 (Mexico) ... 149
F 1–6. Badhamia spinispora (AH 42974): 1. Plasmodiocarps. 5–6. Spores (SEM). Cribraria
confusa (AH 42976): 2–4. Sporocarps. Bars: 1 = 1 mm; 2–4 = 0.5 mm; 5–6 = 2 µm.
S : MEXICO. S: Loc. 4, on bark of Forchhammeria watsonii,
leg. AG & AS, 27-V-2009, obtained from moist chamber, 30-VI-2009 (UES 8129, AH
42975).
C—Comatricha mirabilis is characterized by its capillitium that
extends parallel along the columella without forming a network with bifurcated
ends that protrude from the sporotheca and fully reticulate globose spores. It
was described growing over goat manure in USA in a moist chamber (Benjamin
& Poitras 1950). Later, Mandell & Blackwell (2008) gathered abundant material
from tree barks of Tamarix arabica, Prosopis juliora, Phoenix dactylifera, and
Cordia myxa from a Bahrain desert in the Kingdom of Bahrain archipelago
near Saudi Arabia, which they developed in a moist chamber. Andrade-Bezerra
et al. (2008) described material developed on Brazilian cottontail (Sylvilagus
brasiliensis L.) dung in a moist chamber, Poulain et al. (2011) gathered it from
France on wood debris, and Schnittler et al. (2013) identied the species from
arid areas in China. We obtained three sporocarps on Forchhammeria watsonii
bark in a moist chamber. is is the rst record for Mexico.
150 ... Lizárraga & al.
Comatricha tenerrima (M.A. Curtis) G. Lister
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AS & AG, 22-V-
2009, obtained from moist chamber, 15-VI-2009 (UES 8124); Loc. 9, on decomposing
wood, leg. CS, AS & AG, 6-XI-2009, obtained from moist chamber, 11-XII-2009 (UES
8685); Loc. 10, on decomposing wood, leg. AG & AS, 21-V-2009, obtained from moist
chamber, 6-VI-2009 (UES 8130).
C— Lizárraga et al. (2007) published a description with SEM
micrographs of C. tenerrima from Sonora. It has also been cited from other
natural protected areas in Sonora (Lizárraga et al. 2008a,b; Esqueda et al. 2013).
*Cribraria confusa Nann.-Bremek. & Y. Yamam. F. 2–4
Fructications small (0.3–0.9 mm high), stipitate. Sporotheca globose to
subglobose, 0.1–0.2 mm diam. Stipe 0.2–0.7 mm long, dark brown, with debris.
Peridium evanescent, forming a small discoid calyculus at the base beneath
an open network of irregular attened nodules covered by dictydine granules.
Hypothallus membranous. Spores 7–8 µm diam., yellow in mass, light yellow
under LM, globose, with a warty ornamentation.
S : MEXICO. S: Loc. 3, on palm bers, leg. ML, CS, AS &
AG, 19-VIII-2009, obtained from moist chamber, 5-IX-2009 (UES 8570); obtained
from moist chamber, 7-IX-2009 (UES 8759, AH 42976 with Arcyria cinerea and
Paradiacheopsis sp.); on Guaiacum coulteri A. Gray, obtained from moist chamber, 3-X-
2009 (UES 8571).
C—Cribraria confusa has been recorded only from Chihuahua on
decomposing wood of Pinus sp. by Lizárraga et al. (2003), who presented LM
and SEM micrographs.
Cribraria violacea Rex
S : MEXICO. S: Loc. 10, on palm bers, leg. AG, 21-V-2009,
obtained from moist chamber, 15-VI-2009 (UES 8135).
C— Moreno et al. (2001) published an SEM study of the spore
ornamentation on material from Baja California.
Previously cited from Sonora by Moreno et al. (2006) and Lizárraga et al.
(2008b).
Dictydiaethalium plumbeum (Schumach.) Rostaf.
S : MEXICO. S: Loc. 1, on decomposing wood, leg. ML, CS, AS
& AG, 17-VIII-2009, obtained from moist chamber, 9-IX-2009 (UES 8572); Loc. 5, leg.
AG & AS, 14-III-2009, obtained from moist chambers, 1-V-2009 (UES 7880).
C—Previously cited from Sonora by Moreno et al. (2006) and
Lizárraga et al. (2008b).
Myxomycetes of Sonora 6 (Mexico) ... 151
F –. Comatricha mirabilis (AH 42975): 7. Sporocarp. 8. Capillitium. 9. Spores.
10. Capillitium endings (detail, SEM). 11–12. Spores by SEM. Didymium eremophilum (AH
42977): 13. Sporocarps. 14. Sporocarp (detail). 15–17. Variability of spore ornamentation (SEM).
Bars: 7 = 0.1 mm; 8 = 50 µm; 9 = 10 µm; 13–14 = 0.5 mm; 15 = 5 µm; 16–17 = 2 µm.
152 ... Lizárraga & al.
Didymium dubium Rostaf.
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AS & AG,
9-III-2009, obtained from moist chamber, 20-IV-2009 (UES 7888); Loc. 2, on Prosopis
glandulosa, leg. ML, CS, AS & AG, 19-VIII-2009, obtained from moist chamber, 9-IX-
2009 (UES 8755); Loc. 5, on Prosopis glandulosa, leg. AS & AG, 14-III-2009, obtained
from moist chamber, 9-IV-2009 (UES 7886); Loc. 6, on Prosopis glandulosa, leg. AG
& AS 12-III-2009, obtained from moist chamber, 25-IV-2009 (UES 7889); Loc. 8, on
Cylindropuntia sp., leg. AG & AS, 14-III-2009, obtained from moist chamber, 14-IV-
2009 (UES 7882); Loc. 9, on Cylindropuntia arbuscula, leg. CS, AS & AG, 6-XI-2009,
obtained from moist chamber, 26-XI-2009 (UES 8692).
C—Moreno et al. (2001) published a SEM study of the spore
ornamentation. Previously cited from Sonora by Moreno et al. (2006).
*Didymium eremophilum M. Blackw. & Gilb. F 13–17
Fructications stipitate, gregarious, 0.2–0.9 mm tall. Sporotheca globose to
subglobose, 0.1–0.3 mm diam., whitish with lighter zones, with a dark disc
at the base. Stipe robust, 0.1–0.5 mm tall, reddish-brown to darkish brown,
wider and darker at the base. Hypothallus membranous, discoid, concolorous
with stipe. Peridium double, with the outer membrane formed by calcareous
crosshairs, the inner membranous with iridescent blue tones. Capillitium and
columella absent. Spores globose to subglobose, black in mass, violaceous
by LM, 9–10 µm diam., with a reticulate hemisphere and the opposite warty
(as conrmed by SEM; see Moreno & Mitchell 2013).
S : MEXICO. S: Loc. 4, leg. CS, AS & AG, 7-IX-2009, obtained
from moist chamber, 4-XII-2009 (UES 8698); Loc. 8, on Lophocereus schottii, leg. AG
& AS, 21-V-2009, obtained from moist chamber, 11-VI-2009 (UES 8141, AH 42977);
obtained from moist chamber, 15-VI-2009 (UES 8142); obtained from moist chamber,
19-VI-2009 (UES 8143); obtained from moist chamber, 22-VI-2009 (UES 8144);
obtained from moist chamber, 30-VI-2009 (UES 8145); leg. CS, AS & AG, 6-XI-2009,
obtained from moist chamber, 30-XI-2009 (UES 8699); obtained from moist chamber,
11-XII-2009 (UES 8700); obtained from moist chamber, 12-XII-2009 (UES 8701);
obtained from moist chamber, 12-XII-2009 (UES 8702); Loc. 9, on cactus remains, leg.
AG & AS, 26-V-2009, obtained from moist chamber, 15-VI-2009 (UES 8146).
C—Didymium eremophilum is a succulenticolous species adapted to
arid environments (Blackwell & Gilbertson 1980b), characterized by its small
size, stipitate sporocarps, crystalline lime, lack of capillitium, partly reticulate
verrucose spores, and succulenticolous habitat (Moreno & Mitchell 2013).
Didymium eremophilum specimens, very dicult to nd due to their small
size, have thus far been found only from moist chamber. Previously recorded in
Mexico only from Puebla (Estrada-Torres et al. 2009), our study provides a rst
record for the Sonoran mycobiota.
Myxomycetes of Sonora 6 (Mexico) ... 153
Didymium squamulosum (Alb. & Schwein.) Fr.
S : MEXICO. S: Loc. 4, on deer manure, leg. AG & AS 12-
III-2009, obtained from moist chamber, 1-V-2009 (UES 7881); Loc. 7, on Stenocereus
thurberi, leg. AG & AS, 11-III-2009, obtained from moist chamber, 6-IV-2009 (UES
7890).
C—Lizárraga et al. (1999a) published a SEM study of the spore
ornamentation. Widely cited for Mexico (Moreno et al. 2007), D. squamulosum
has been previously cited for Sonora by Moreno et al. (2006) and Lizárraga et
al. (2008b).
Didymium sturgisii Hagelst. F 18–23
S : MEXICO. S: Loc. 7, on Prosopis glandulosa, leg. AG & AS,
11-III-2009, obtained from moist chamber, 8-IV-2009 (UES 7884, AH 42978).
C—is species is characterized by its dirty white to whitish grey
attened to subglobose fructications (occasionally with small supercial
orices) and sporocarp interiors with erect pillars or internally traversing
trabeculae. Moreno et al. (2001) published an SEM study of the capillitium,
pillars, and spores.
Didymium sturgisii has been recorded from Baja California (Moreno et al.
2007), Chihuahua (Salazar-Márquez et al. 2014), Puebla (Estrada-Torres et al.
2009), and Sonora (Esqueda et al. 2012, 2013).
Echinostelium apitectum K.D. Whitney
S : MEXICO. S: Loc. 1, on Prosopis glandulosa leg. AS & AG,
9-III-2009, observed in moist chamber, 28-III-2009, 4-IV-2009 (UES 7892); Loc. 3, on
Stenocereus thurberi, 19-VIII-2009, obtained in moist chamber, 2-IX-2009, 5-IX-2009
(UES 8589); Loc. 4, on Olneya tesota, leg. AG & AS, 3-VI-2009, obtained from moist
chamber, 17-VI-2009 (UES 8149); Loc. 5, on decomposing wood, leg. ML, CS, AS &
AG, 18-VIII-2009, obtained in moist chamber, 2-IX-2009, 5-IX-2009 (UES 8581); Loc.
6, on Prosopis glandulosa, leg. AG & AS, 12-III-2009, observed in moist chamber, 28-
III-2009, 2-IV-2009 (UES 7902); Loc. 7, on Prosopis glandulosa, leg. AG & AS, 22-V-
2009, observed in moist chamber, 3-VI-2009, 22-VI-2009 (UES 8151); Loc. 9, Carnegiea
gigantea, leg. ML, CS, AS & AG, 20-VIII-2009, observed in moist chamber, 2-IX-2009,
7-IX-2009 (UES 8584); Loc. 10, on Prosopis glandulosa, leg. CS, AS & AG, 3-XI-2009,
observed in moist chamber, 9-XI-2009, 30-XI-2009 (UES 8703).
C—Previously cited from Sonora by Lizárraga et al. (2007, 2008a).
Echinostelium arboreum H.W. Keller & T.E. Brooks
S : MEXICO. S: Loc. 1, on decomposing wood, leg. AG & AS,
22-V-2009, obtained from moist chamber, 11-VI-2009 (UES 8106 with Arcyria cinerea).
154 ... Lizárraga & al.
F –. Didymium sturgisii (AH 42978): 18. Fructications attened with some calcium
carbonate granules. 19. Sporocarp (detail of small supercial orices). 20. Capillitium and
erect pillars. 21–22. Spores (SEM). 23. Spore ornamentation (detail, SEM). Licea denudescens
(AH 42979): 24. Sporocarps. 25. Spore (SEM). Licea succulenticola (AH 42980): 26. Sporocarps.
27. Spore (SEM). Bars: 18–19 = 0.5 mm; 20 = 0.25 mm; 21–22 = 2 µm; 23 = 1 µm.
C—Moreno et al. (2001) published a SEM study of the sporocarp and
spores. Previously cited from Sonora by Esqueda et al. (2013), E. arboreum has
been widely reported from Mexico (Moreno et al. 2007).
Echinostelium colliculosum K.D. Whitney & H.W. Keller
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AS & AG,
9-III-2009, 28-III-2009, obtained from moist chamber, 4-IV-2009 (UES 7891); Loc.
5, on Guaiacum coulteri, 14-III-2009, obtained from moist chamber, 4-IV-2009 (UES
7893); on Olneya tesota, observed in moist chamber, 4-IV-2009 (UES 7894); on Prosopis
glandulosa, obtained from moist chamber, 8-IV-2009 (UES 7895); on Parkinsonia sp.,
leg. AG & AS, 26-V-2009, obtained from moist chamber, 11-VI-2009 (UES 8152); Loc.
6, on Cylindropuntia arbuscula, leg. AS & AG, 27-V-2009, 3-VI-2009, obtained from
moist chamber, 11-VI-2009 (UES 8153); on Parkinsonia sp., leg. ML, CS, AS & AG, 19-
VIII-2009, in moist chamber, 2-IX-2009, 5-IX-2009 (UES 8588); Loc. 7, on Parkinsonia
sp., leg. AS & AG, 11-III-2009, in moist chamber, 9-IV-2009 (UES 7901); on Encelia
farinosa A. Gray ex Torr., 22-V-2009, 3-VI-2009, in moist chamber, 11-VI-2009 (UES
8150); on decomposing wood, leg. ML, CS, AS & AG, 17-VIII-2009, in moist chamber,
2-IX-2009, 5-IX-2009 (UES 8591); Loc. 9, on Guaiacum coulteri, 11-III-2009, 28-III-
2009, in moist chamber, 4-IV-2009 (UES 7896); on Olneya tesota, in moist chamber,
4-IV-2009 (UES 7898); on Cylindropuntia arbuscula, in moist chamber, 4-IV-2009 (UES
7899); leg. ML, CS, AS & AG, 20-VIII-2009, in moist chamber, 2-IX-2009, 7-IX-2009
(UES 8583); Loc. 10, on Cylindropuntia sp., leg. AS & AG, 12-III-2009, 4-IV-2009 (UES
7900); leg. ML, CS, AS & AG, 18-VIII-2009, in moist chamber, 9-IX-2009 (UES 8582);
Loc. 10, on Cylindropuntia sp., leg. AS & AG, 12-III-2009, 4-IV-2009 (UES 7900).
C—is species is characterized by its (60–)75–90 µm high stipitate
sporocarps, globose 9–13 µm diam. pale yellow spores that have a spine-like
ornamentation and articular spore wall. A LM study of sporocarps and spores
by Lizárraga et al. (2003) and SEM study by Moreno et al. (2001) have been
published.
Echinostelium colliculosum has been recorded for Baja California Sur and
Chihuahua (Moreno et al. 2007), Oaxaca and Puebla (Estrada-Torres et al.
2009), and Sonora (Esqueda et al. 2013).
Echinostelium minutum de Bary
S : MEXICO. S: Loc. 1, on Prosopis glandulosa, leg. ML, CS, AS
& AG, 17-VIII-2009, in moist chamber, 2-IX-2009, 7-IX-2009 (UES 8580); Loc. 3, on
Stenocereus thurberi, leg. ML, CS, AS & AG, 17-VIII-2009, 7-IX-2009 (UES 8590); Loc. 7,
on Prosopis glandulosa, leg. AG, 22-05-2009, obtained from moist chamber, 11-06-2009
Myxomycetes of Sonora 6 (Mexico) ... 155
156 ... Lizárraga & al.
(UES 8154)
; Loc. 9, on Carnegiea gigantea, 9-IX-2009 (UES 8585); Loc. 10, on Prosopis
glandulosa, leg. CS, AS & AG, 3-XI-2009, in moist chamber, 9-XI-2009, II-2009 (UES
8704).
C— Moreno et al. (2001) published a SEM study of the sporocarp
and spores. Previously reported from Sonora by Moreno et al. (2006).
Enerthenema papillatum (Pers.) Rostaf.
S : MEXICO. S: Loc. 4, on Forchhammeria watsonii, leg. ML,
CS, AS & AG, 19-VIII-2009, obtained in moist chamber, 5-IX-2009 (UES 8758).
C—Lizárraga et al. (1999a) published a SEM study of the spores from
collections from Baja California, Mexico. Widely recorded in Mexico (Moreno
et al. 2007), E. papillatus was recorded previously from Sonora by Moreno et
al. (2006).
Fuligo cinerea (Schwein.) Morgan
S : MEXICO. S: Loc.10, on sheep manure, leg., AG, 21-V-2009,
obtained from moist chamber, 30-VI-20009 (UES 8118).
C—Sometimes similar in size as some Badhamia species (e.g.,
B. spinispors, B. verrucospora G. Moreno et al.), Fuligo cinerea is separated by
its ovoid spores covered by small warts that join together to form a reticulum.
Cited previously for Sonora by Braun & Keller (1986) and Lizárraga et al.
(2007).
Fuligo septica (L.) F.H. Wigg.
S : MEXICO. S: Loc. 9, on soil, leg. ML, 11-III-2009 (UES
8677).
C—Lizárraga et al. (1999a) published a SEM study of the spores.
Fuligo septica has a wide distribution in Mexico (Moreno et al. 2007) and has
been cited for Sonora by Pérez-Silva et al. (2001) and Lizárraga et al. (2007,
2008a,b).
Licea biforis Morgan
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AG & AS, 9-III-
2009, obtained in moist chamber, 6-IV-2009 (UES 7905); Loc. 2, on Prosopis glandulosa,
leg. AS & AG, 14-III-2009, obtained in moist chamber, 13-IV-2009 (UES 7908); Loc. 3,
on Acacia sp., 26-V-2009, obtained in moist chamber, 15-VI-2009 (UES 8159); Loc. 5,
on Guaiacum coulteri bark, leg. AS & AG 14-III-2009, obtained in moist chamber, 20-
IV-2009 (UES 7913); Loc. 6, on Prosopis glandulosa, leg. AG & AS, 27-V-2009, obtained
in moist chamber, 15-VI-2009 (UES 8162); Loc. 9, on Olneya tesota, leg. AG & AS, 11-
III-2009, obtained in moist chamber, 1-V-2009 (UES 7915); Loc. 10, leg. CS, AS & AG,
3-XI-2009, obtained in moist chamber, 12-XII-2009 (UES 8716).
Myxomycetes of Sonora 6 (Mexico) ... 157
C—Lizárraga et al. (1998, 1999b) published SEM studies of the
spores. Widely distributed in Mexico (Moreno et al. 2007), Licea biforis was
previously cited from Sonora by Moreno et al. (2006).
Licea denudescens H.W. Keller & T.E. Brooks F 24–25
Fructications sessile, globose to subglobose, 0.1–0.3 mm diam., 0.1 mm
height, brown to blackish. Peridium membranous with some iridescent tones,
internally ornamented with small buds. Spores violaceous in mass, yellowish
brown by LM, 10–11 µm diam., globose to subglobose, walls thick with pale
(thinner) areas, smooth by SEM.
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AS & AG, 9-III-
2009, obtained in moist chamber, 8-IV-2009 (UES 7916, AH 42979); Loc. 5, on Olneya
tesota, leg. CS, AS & AG, 6-XI-2009, obtained in moist chamber, 12-XII-2009 (UES
8711); Loc. 7, on Parkinsonia sp., 11-III-2009, obtained in moist chamber, 20-IV-2009
(UES 7918); obtained in moist chamber, 1-V-2009 (UES 7917); Loc. 8, on Acacia greggii
A. Gray, leg. ML, CS, AS & AG, 18-VIII-2009, obtained in moist chamber, 3-X-2009
(UES 8605); Loc. 9, on cactus remains, 11-III-2009, observed in moist chamber, 15-IV-
2009 (UES 7919).
C—Licea denudescens is diagnosed by its thick-walled spores with
thinner areas and papillae found on the inside of the peridium. It has been
cited in Mexico from Puebla (Estrada-Torres et al. 2009), Chihuahua (Salazar-
Márquez et al. 2014), and Sonora (Esqueda et al. 2013).
Licea kleistobolus G.W. Martin
S : MEXICO. S: Loc. 8, on Acacia greggii bark, leg. ML, CS, AS
& AG, 18-VIII-2009, observed in moist chamber, 7-IX-2009 (UES 8608).
C—Moreno et al. (2001) published a SEM study of Licea kleistobolus,
which is characterized by its circular to subglobose fructications with a
characteristic operculum and warted spores with groups of denser warts. Cited
previously from Sonora by Moreno et al. (2006) and Lizárraga et al. (2007).
Licea succulenticola Mosquera, Lado, Estrada & Beltrán-Tej. F 26–27
Fructications sessile, yellowish to dark-yellow at mature, dispersed to
gregarious, subglobose, oval to elongated, 0.1–0.3 mm long, 0.1–0.2 mm
diam., and 0.1 mm tall, with an apical and longitudinal dehiscence. Peridium
membranous, ornamented on its inner part with small warts. Spores yellow in
mass, light yellow by LM, globose to subglobose, (11–)12–14 µm diam., with
a lightly paler area, with a slightly warty ornamentation. SEM examination
shows that the spore ornamentation is formed by small croziers more or less
uniformly distributed over the in spore surface.
S : MEXICO. S: Loc. 2, on Opuntia sp., 5-XI-2009, observed in
moist chamber, 14-XII-2009 (UES 8712); Loc. 3, on Cylindropuntia sp., leg. AS & AG,
158 ... Lizárraga & al.
14-III-2009, obtained in moist chamber, 28-III-2009, 20-IV-2009 (UES 7924); obtained
in moist chamber, 25-IV-2009 (UES 7925, 7926); Loc. 4, on Lophocereus schottii, leg.
AS & AG, 12-III-2009, obtained from moist chamber, 20-IV-2009 (UES 7923); Loc.
6, on Lophocereus schottii, leg. CS, AS & AG, 7-XI-2009, observed in moist chamber,
9-XI-2009, 9-XII-2009 (UES 8726); Loc. 7, on Lophocereus schottii, leg. AS & AG, 11-
III-2009, observed in moist chamber, 28-III-2009, 20-IV-2009 (UES 7927); obtained in
moist chamber, 1-V-2009 (UES 7928, 7929); on Stenocereus thurberi, leg. ML, CS, AS &
AG, 17-VIII-2009, observed in moist chamber, 2-IX-2009, 7-IX-2009 (UES 8747, AH
42980); obtained in moist chamber, 14-IX-2009 (UES 8602); obtained in moist chamber,
21-IX-2009 (UES 8604); obtained in moist chamber, 21-IX-2009 (UES 8603); Loc. 8, on
Lophocereus schottii, leg. ML, CS, AS & AG, 18-VIII-2009, observed in moist chamber,
2-IX-2009, 9-IX-2009 (UES 8594); obtained in moist chamber, 14-IX-2009 (UES 8595);
obtained in moist chamber, 24-IX-2009 (UES 8596); obtained in moist chamber, 30-
IX-2009 (UES 8597); obtained in moist chamber, 5-X-2009 (UES 8598, AH 42981); on
Stenocereus thurberi, leg. CS, AS & AG, 6-XI-2009, observed in moist chamber, 9 -XI-
2009, 14-XII-2009 (UES 8725); Loc. 10, on Cylindropuntia arbuscula, leg. AG & AS,
12-III-2009, obtained in moist chamber, 20-IV-2009 (UES 7920); on cactus remains,
obtained in moist chamber, 20-IV-2009 (UES 7921); obtained in moist chamber, 1-V-
2009 (UES 7922); on Lophocereus schottii, leg. ML, CS, AS & AG, 18-VIII-2009, obtained
in moist chamber, 2-IX-2009, 17-IX-2009 (UES 8599); obtained in moist chamber, 30-
IX-2009 (UES 8600); obtained in moist chamber, 30-IX-2009 (UES 8601).
C—Licea succulenticola resembles L. pumila G.W. Martin & R.M.
Allen (Martin & Alexopoulos 1969) in sporothecal form and spore colour.
(see Mosquera et al. 2003 for dierences). Previously recorded from Hidalgo,
Morelos, Puebla, and Tlaxcala (Mosquera et al. 2003), Puebla (Estrada-Torres
et al. 2009), Sonora (Esqueda et al. 2013), and Chihuahua (Salazar-Márquez et
al. 2014).
Paradiacheopsis mbriata (G. Lister & Cran) Hertel ex Nann.-Bremek. F 28–30
S : MEXICO. S: Loc. 3, on Cylindropuntia sp., leg. AG,
14-III-2009, observed in moist chamber, 28-III-2009, 13-IV-2009 (UES 7943,
AH 42982).
C—Widely distributed in Mexico (Moreno et al. 2007), P. mbriata
was reported from Sonora by Moreno et al. (2006).
Perichaena depressa Lib.
S : MEXICO. S: Loc. 1, on rabbit manure, leg. ML, CS, AS &
AG, 17-VIII-2009, obtained in moist chamber, 9-X-2009 (UES 8628); Loc. 9, on horse
manure, 20-VIII-2009, obtained in moist chamber, 5-X-2009 (UES 8629).
C—Lizárraga et al. (1998) and Moreno et al. (2001) published SEM
studies of spores and capillitium, comparing type material with samples from
Baja California and Sinaloa. Widely distributed throughout Mexico (Moreno
et al. 2007), P. depressa was previously cited for Sonora by Moreno et al. (2006).
Myxomycetes of Sonora 6 (Mexico) ... 159
F –. Paradiacheopsis mbriata (AH 42982): 28. Sporocarp. 29. Sporotheca (detail).
30. Fiber stalk (detail). Perichaena stipitata (AH 42983): 31–32. Stipitate sporocarp (detail).
33. Peridial inner surface (detail). 34. Capillitium with irregular outline. 35–36. Spores (SEM).
Bars: 28–29 = 0.1 mm; 30 = 0.02 mm; 31–32 = 0.5 mm; 33, 35–36 = 1 µm; 34 = 5 µm.
*Perichaena luteola (Kowalski) Gilert
S : MEXICO. S: Loc. 10, on cow manure, leg. ML, CS, AS &
AG, 18-VIII-2009, obtained in moist chamber, 27-IX-2009 (UES 8749); 3-X-2009 (UES
8627).
C—e above collections have 10–13 µm diam. globose to
subglobose spores that are yellow in mass (lighter yellow by LM) and with a
verruculose episporium and a yellow 2–3µm diam. capillitium with smooth
surface. Lizárraga et al. (1999b) published a SEM study of material from Baja
California. Perichaena luteola is widely distributed in Mexico (Moreno et al.
2007).
160 ... Lizárraga & al.
*Perichaena stipitata Lado, Estrada & D. Wrigley F 31–36
Fructications short-stipitate, solitary to gregarious, 0.3–0.8 mm tall. Stipe
robust, cylindrical, black, 0.1–0.2 mm long. Sporotheca globose to subglobose,
rarely obpyriform, yellow, 0.2–0.5 mm diam. Peridium brownish on the base,
inner surface ornamented with small rings by LM and SEM. Hypothallus dark,
membranous and irregular. Capillitium sparse, formed by yellowish branched
laments of approximately 2 µm diam., exible, of irregular outline. Spores
yellow in mass, light yellow by LM, 12–15 µm diam., globose, warted. SEM
examination shows that the spore is ornamented by warts with smooth apices
and which sometimes join into small groups.
S : MEXICO. S: Loc. 9, on cactus remains, leg. AG & AS, 26-
V-2209, obtained in moist chamber, 30-VI-2009 (UES 8210, AH 42983); on Lophocereus
schottii, leg. ML, CS, AS & AG, 20-VIII-2009, obtained in moist chamber, 17-IX-2009
(UES 8762); obtained in moist chamber, 24-IX-2009 (UES 8761); obtained in moist
chamber, 3-X-2009 (UES 8760, AH 42984); obtained in moist chamber, 9-X-2009 (UES
8643); Loc. 10, on Lophocereus schottii, leg. ML, CS, AS & AG, 18-VIII-2009, obtained
in moist chamber, 5-X-2009 (UES 8642).
C—Stipitate fructications and an inner peridium ornamented by
small rings characterize this succulenticolous species. First described as new
to science by Estrada-Torres et al. (2009) from Puebla and Queretaro, Mexico,
P. stipitata was later described from Arizona (USA) by Moreno & Mitchell (2013),
who published a colour plate showing its development and morphological
characteristics. A new record for Sonora.
Perichaena vermicularis (Schwein.) Rostaf.
S : MEXICO. S: Loc. 1, on Carnegiea gigantea, leg. CS, AS
& AG, 5-XI-2009, obtained in moist chamber, 11-XII-2009 (UES 8734); Loc. 2, on
Cylindropuntia arbuscula, leg. AS & AG, 22-V-2009, obtained in moist chamber, 25-VI-
2009 (UES 8191); Loc. 3, on Cylindropuntia sp., leg. AS & AG 14-III-2009, obtained in
moist chamber, 15-IV-2009 (UES 7953); Loc. 4, on Forchhammeria watsonii, leg. AG &
AS, 12-III-2009, obtained in moist chamber, 3-IV-2009 (UES 7944); Loc. 6, Parkinsonia
sp., 19-VIII-2009, obtained in moist chamber, 24-IX-2009 (UES 8633); Loc. 7, on
Cactaceae remains, 11-III-2009, obtained in moist chamber, 8-IV-2009 (UES 7947);
obtained in moist chamber, 20-IV-2009 (UES 7948); Loc. 8, on Cactaceae remains,
obtained in moist chamber, 18-VIII-2009, 17-IX-2009 (UES 8631).
C—Moreno et al. (2001) published a SEM study of spores and
capillitium in collections from Baja California that grew cespitose in moist
chamber over dierent substrates. Widely distributed in Mexico (Moreno et
al. 2007), P. vermicularis was previously reported from Sonora by Moreno et
al. (2006).
Myxomycetes of Sonora 6 (Mexico) ... 161
Physarum album (Bull.) Chevall.
S : MEXICO. S: Loc. 7, on Prosopis glandulosa, leg. AG & AS,
17-VIII-2009, obtained in moist chamber, 9-IX-2009 (UES 8651); Loc. 10, on Prosopis
glandulosa, leg. ML, CS, AS & AG, 18-VIII-2009, obtained in moist chamber, 7-IX-2009
(UES 8650).
C—Lizárraga et al. (1999b) published a SEM study of spore
ornamentation in collections from Baja California. Widely distributed in
Mexico (Moreno et al. 2007) and previously reported for Sonora (Lizárraga
et al. 2007, 2008b), Physarum album frequently grows on many dierent
substrates when cultivated in moist chamber.
Physarum decipiens M.A. Curtis
Fructications sessile, dispersed to grouped, subglobose or forming
short plasmodiocarp. Sporotheca yellow-greenish, 0.3–1.2 × 0.2–0.6 mm,
membranous, peridium encrusted with calcium carbonate. Capillitium
physaroid. Spores 11–13 µm diam., subglobose to globose, violaceous and
slightly warty.
S : MEXICO. S: Loc. 1, on Prosopis glandulosa, leg. AG & AS,
9-III-2009, obtained in moist chamber, 6-IV-2009 (UES 7967); 22-V-2009, obtained in
moist chamber, 11-VI-2009 (UES 8193); L oc. 2, on Prosopis glandulosa, leg. AS & AG 14-
III-2009, obtained in moist chamber, 8-IV-2009 (UES 7970); Loc. 4, on Forchhammeria
watsonii, leg. ML, CS, AS & AG, 19-VIII-2009, obtained in moist chamber, 7-IX-2009
(UES 8649).
C—Morphologically, P. decipiens is highly variable. Previously cited
from Chihuahua (Lizárraga et al. 2003), Oaxaca and Puebla (Estrada-Torres et
al. 2009), Tlaxcala (Rodríguez-Palma et al. 2002), and recently Sonora (Esqueda
et al. 2012, 2013).
Physarum leucophaeum Fr.
S : MEXICO. S: Loc. 1, on Olneya tesota, leg. AS & AG, 22-V-
2009, obtained in moist chamber, 25-VI-2009 (UES 8194); Loc. 5, on Parkinsonia sp.,
leg. AS & AG 14-III-2009, obtained in moist chamber, 1-V-2009 (UES 7966); Loc. 6, on
Encelia farinosa, 27-V-2009, obtained in moist chamber, 22-VI-2009 (UES 8198); Loc.
8, on Larrea tridentata Coville, leg. AS & AG, 21-V-2009, obtained in moist chamber,
30-VI-2009 (UES 8202); Loc. 10, on Guaiacum coulteri, leg. ML, CS, AS & AG, 18-VIII-
2009, obtained in moist chamber, 9-X-2009 (UES 8653).
C— Moreno et al. (2001) published a SEM study of spore
ornamentation in collections from Baja California. e recently named
species from arid regions of Eurasia and North and South America, Physarum
pseudonotabile Novozh. et al. (Novozhilov et al. 2013), is very close to P. notabile
162 ... Lizárraga & al.
T. Macbr. and P. leucophaeum; distinguishing the three species is complex
and additional ecological, morphological, and molecular studies are needed.
Physarum leucophaeum is widely distributed in Mexico (Moreno et al. 2007),
common when cultured in moist chamber. Previously cited from Sonora by
Lizárraga et al. (2007, 2008b).
Acknowledgments
is research was made possible by support from the following projects: CONABIO
(Proyecto GT016). We wish to express our gratitude to Dr. S.L. Stephenson (University
of Arkansas, U.S.A.) and Mr. D.W. Mitchell (Upper Harteld, U.K.) for reviewing the
manuscript and providing useful comments. We also extend our thanks to Mr. A. Priego
and Mr. J.A. Pérez (Electron Microscopy Service, University of Alcalá de Henares)
for their invaluable help with the SEM. We also thank Luis Monje (Department of
Drawing and Scientic Photography, University of Alcalá de Henares) for his help in
digitally preparing the photographs, and we are grateful to Dr. J. Rejos, curator of the
AH herbarium for his assistance with the specimens examined in the present study.
M. Lizárraga extends his gratitude to Dr. Alejandro Martínez Martínez (Jefe del Dpto.
de Ciencias Químico Biológicas, ICB-UACJ) for his assistance in obtaining nancial
support. C. Salazar thanks CONACYT for her scholarship to obtain a master’s degree.
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