A review of the taxonomy and species diversity in Dacrymycetes (Fungi, Basidiomycota) in Mexico

Abstract

The class Dacrymycetes has been considered a monophyletic group based on shared morphological characteristics and recent molecular studies, with bifurcate basidia and non perforate parenthosome as synapomorphies. In this review of the state of knowledge for Dacrymycetes (dacrymycetoid fungi) in Mexico, a detailed survey of the literature revealed 21 species in eight genera in Mexico; 20 belong to seven genera of Dacrymycetaceae and one species belongs to the Cerinomycetaceae. The best-represented genus is Dacrymyces, and the most-recorded species is Dacryopinax spathularia. Taxa of Dacrymycetes in Mexico have been recorded from 22 states, mainly those located in central Mexico. The members of this group recorded in Mexico are, apparently, wood-decomposers and have been most frequently collected during the wet season (end of May to beginning of October), but they can be found throughout the year. From the review, it is evident that systematic and biogeographic knowledge on the Dacrymycetes in Mexico is scarce. Thus, more extensive sampling and detailed systematic and molecular studies are needed.

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© 2017 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, www.borntraeger-cramer.de
Germany. DOI: 10.1127/nova_hedwigia/2017/0417 0029-5035/2017/0417 $ 5.00
Nova Hedwigia Vol. 105 (2017) Issue 3–4, 365–384
published online April 24, 2017; published in print November 2017 Article
C
A review of the taxonomy and species diversity in
Dacrymycetes (Fungi, Basidiomycota) in Mexico
Sandra Castro-Santiuste1, Sigfrido Sierra2, Laura Guzmán-Dávalos3
and Isolda Luna-Vega1*
1 Laboratorio de Biogeografía y Sistemática, Departamento de Biología Evolutiva,
Facultad de Ciencias, Universidad Nacional Autónoma de México, Ciudad
Universitaria, Av. Universidad 3000, Coyoacán, 04510, Ciudad de México, Mexico
2 Laboratorio de Taxonomía de Hongos Tremeloides (Heterobasidiomycetes),
Departamento de Biología Comparada, Facultad de Ciencias, Universidad Nacional
Autónoma de México, Ciudad Universitaria, Av. Universidad 3000, Coyoacán, 04510,
Ciudad de México, Mexico
3 Laboratorio de Micología, Departamento de Botánica y Zoología, Universidad de
Guadalajara, Apdo. Postal 1–131, 45101, Zapopan, Jal., Mexico
With 4 gures and 2 tables
Abstract: The class Dacrymycetes has been considered a monophyletic group based on shared
morphological characteristics and recent molecular studies, with bifurcate basidia and non perforate
parenthosome as synapomorphies. In this review of the state of knowledge for Dacrymycetes
(dacrymycetoid fungi) in Mexico, a detailed survey of the literature revealed 21 species in eight
genera in Mexico; 20 belong to seven genera of Dacrymycetaceae and one species belongs to the
Cerinomycetaceae. The best-represented genus is Dacrymyces, and the most-recorded species is
Dacryopinax spathularia. Taxa of Dacrymycetes in Mexico have been recorded from 22 states, mainly
those located in central Mexico. The members of this group recorded in Mexico are, apparently,
wood-decomposers and have been most frequently collected during the wet season (end of May to
beginning of October), but they can be found throughout the year. From the review, it is evident that
systematic and biogeographic knowledge on the Dacrymycetes in Mexico is scarce. Thus, more
extensive sampling and detailed systematic and molecular studies are needed.
Key words: bifurcate basidia, Dacrymyces, Dacryopinax, distribution, saprobic fungi.
*Corresponding author: luna.isolda@gmail.com
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Introduction
The Dacrymycetes, Tremellomycetes and two orders of Agaricomycetes (Auri-
culariales and part of Sebacinales), commonly known as jelly fungi, are highly
diverse but inconspicuous macrofungi. Their classification and species and genera
delimitation have been problematic because of the morphological heterogeneity of
their basidia, basidiospore germination, and basidiome form (Sierra & Cifuentes
1993, Oberwinkler 2014). Although those in the class Agaricomycetes mainly have
a single uniform type of basidia, Auriculariales have transversal septate basidia,
and Sebacinales have longitudinal septate basidia.
Traditional classification schemes, such as that of Martin (1952), suggest that the
basidiomycetes can be segregated into two groups, Heterobasidiomycetes with
septate and aseptate basidia and Homobasidiomycetes with aseptate basidia. Martin
(1952) included the Tremellales within the Heterobasidiomycetes and recognized
ten families within the Tremellales: Auriculariaceae, Hyaloriaceae, Phleogenaceae,
Septobasidiaceae, Sirobasidiaceae, Sporobolomycetaceae, and Tremellaceae with
septate basidia and Ceratobasidiaceae, Dacrymycetaceae, and Tulasnellaceae with
aseptate basidia. From these groups, dacrymycetoid fungi (or Dacrymycetaceae
sensu Martin [1952], Dacrymycetidae sensu Jülich [1981], or Dacrymycetes sensu
Doweld [2001]), have slightly clavate basidia that expand apically to produce two
long, cylindrical sterigmata, so the final shape is like a tuning fork (Martin 1952,
Oberwinkler 2014). Talbot (1954, 1968, 1973) and Donk (1958) reclassified the
basidiomycetes based on the type and structure of the basidia and recognized three
classes: Teliomycetes, Phragmobasidiomycetes, and Holobasidiomycetes. The last
class comprises species with aseptate basidia, including the dacrymycetoid fungi
studied herein.
Recently, the class Dacrymycetes, including the orders Dacrymycetales and
Unilacrymales and families Dacrymycetaceae, Cerinomycetaceae, and Unilacry-
maceae, have been recognized as a natural group (Hibbett 2006, Shirouzu et al. 2013,
Oberwinkler 2014), with the presence of bifurcated holobasidia, with the exception
of Unilacrymales, that have cylindrical holobasidia (Shirouzu et al. 2013) and a
non-perforated parenthosome as two synapomorphic traits (Tehler 1988, Swann &
Taylor 1993, Oberwinkler 1994). Previously, Khan & Kimbrough (1982), using only
morphological characters, highlighted the importance of the septal ultrastructure
for the classification of the basidiomycetes.
The recognition of genera within the group is problematic, reflected by the 16
synonyms found for this taxonomic category. Different authors have considered
between two [Dacrymyces Nees and Calocera (Fr.) Fr., following Neuhoff 1936]
and 12 valid genera (Arrhytidia Berk. & M.A.Curtis, Calocera, Cerinomyces
G.W.Martin, Dacrymyces, Dacryomitra Tul. & C.Tul., Dacryonaema Nannf.,
Dacryopinax G.W.Martin, Ditiola Fr., Femsjonia Fr., Guepiniopsis Pat., Heterotextus
Lloyd, Unilacryma Shirouzu, Tokum. & Oberw., following Oberwinkler 2014
and the present study). On the other hand, classification schemes that rely solely
on morphology of the basidiomata are ambiguous because the basidioma appears
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to be insufficient for generic delimitation. Therefore, other morphological traits
such as micromorphology and ultrastructure need be included to attain a good
generic delimitation (Oberwinkler 2014). Systematic studies aimed at resolving
the taxonomic relationships within the group are few. For example, molecular
studies have cast doubts on the monophyly of Dacrymyces, the type genus of
Dacrymycetes, showing that several of its species are scattered widely throughout
the Dacrymycetales (Shirouzu et al. 2007).
Dacrymycetes has been considered as the sister group to Agaricomycetes according
to Hibbett (2006) and Hibbett et al. (2007). These authors found that Dacrymycetes
nests within the Agaricomycotina, with Agaricomycetes and Tremellomycetes.
The species of Dacrymycetes have a wide range of basidioma types (e.g., resupinate,
pulvinate, turbinate, stipitate, spatulate, and dendroid), mainly with yellow-orange
coloration and a gelatinous to cartilaginous consistency (Oberwinkler 2014). The
vast majority of species generate brown rot and, to a lesser degree, white rot (Seifert
1983). Molecular clock analyses have estimated that Dacrymycetes originated
280–360 million years ago (Dentinger et al. 2010); thus, this group stands as one
of the oldest wood decomposers within the Basidiomycota (Shirouzu et al. 2013).
Oberwinkler (2014) stated that, although molecular phylogenies give support to a
well-defined Dacrymycetes, the taxonomy of the group remains controversial. Thus,
this author proposed a series of detailed systematic studies on the micromorphology
of these species and the inclusion of more molecular markers in phylogenetic
analyses. Such studies have been needed to elucidate the diversity and distribution
of Dacrymycetes groups in Mexico. This study highlights the need for this research
because the actual number of species in Mexico is expected to be greater than
the 21 species presently recorded, especially when compared with the number
recorded for smaller countries such as Japan and Great Britain (Reid 1974: British
Dacrymycetales, 20 species; Shirouzu et al. 2009: Japanese Dacrymycetes 42
species).
As a review of the present state of knowledge on the Dacrymycetes fungi in Mexico,
the present study summarizes for the first time eco-geographical data (e.g., vegetation
type, elevation, geographic distribution), habitat, and phenology of the species
in Mexico. In this context, the present work might serve as a basis for decision-
making regarding the conservation status and management of dacrymycetoid fungi
in Mexico.
Materials and methods
After an exhaustive review of the literature (up to February 2017), a species database containing
389 records was compiled. These records include data on the elevation, geographic coordinates,
vegetation type, locality, habitat, collector, and sampling date of each specimen. From this database,
the number of species per genus was obtained, as well as the eco-geographical distribution and
phenology of the species. Seven historical records are also mentioned. Consulted studies are
shown in Table 1.
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Table 1. Studies that consider Mexican dacrymycetoid fungi through February 2017.
Reference Year
Welden & Lemke 1961
Lowy 1965, 1971, 1980
Guzmán 1972, 1980, 1983, 2003
Mendiola & Guzmán 1973
Welden & Guzmán 1978
Lowy & Guzmán 1979
Varela & Cifuentes 1979
Chio & Guzmán 1982
Frutis & Guzmán 1983
Chacón & Guzmán 1984
Guzmán & Villarreal 1984
Portugal et al. 1985
Díaz-Barriga et al. 1988
Pérez-Moreno & Villarreal 1988
Téllez-Bañuelos et al. 1988
Cifuentes et al. 1990, 1993
Pérez-Silva & Esqueda-Valle 1992
Sierra & Cifuentes 1993, 2005, 2009
Guzmán-Dávalos & Fragoza-Díaz 1995
Moreno-Fuentes et al. 1994, 2004
Esqueda-Valle et al. 1995
Frutis & Pinzón-Picaseño 1997
García-Jiménez et al. 1998
Sierra 2000
Guzmán-Dávalos 2001
López & García 2001, 2002
Herrera-Fonseca et al. 2002
Valenzuela et al. 2004
García-Jiménez & Guevara-Guerrero 2005
García-Jiménez & Valenzuela 2005
Tovar-Velasco & Valenzuela 2006
Marmolejo & Méndez-Cortés 2007
Villarruel-Ordaz & Cifuentes 2007
López-Eustaquio et al. 2010
Rodríguez et al. 2010
Pompa-González et al. 2011
Sánchez-Jácome & Guzmán-Dávalos 2011
Raymundo et al. 2012
Salinas-Salgado et al. 2012
Sierra et al. 2012
Chanona-Gómez et al. 2014
Gándara et al. 2014
Villarruel-Ordaz et al. 2015
Quiñones-Martínez & Garza-Ocañas 2015
Padilla-Velarde et al. 2016
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Results
SpecieS recorded in Mexico: At present, eight genera and 21 species of Dacrymycetes
have been found in Mexico; seven of these genera encompass 20 species belonging to
Dacrymycetaceae, and the other genus is monotypic and belongs to Cerinomycetaceae.
The genus with the greatest number of species in Mexico is Dacrymyces with six species
(Table 2, Fig. 1). Most of the herbarium specimens cited in the literature reviewed
were from ENCB (Escuela Nacional de Ciencias Biológicas, Instituto Politécnico
Nacional, Mexico City) with 133 specimens, followed by FCME (Facultad de Ciencias,
Universidad Nacional Autónoma de México, Mexico City) with 73 specimens, and
IBUG (Instituto de Botánica, Universidad de Guadalajara) with 44 specimens. Herbaria
acronyms are according Thiers (2016).
Arrhytidia was described by Berkeley and Curtis (1849), with A. flava Berk. &
M.A.Curtis as type species from material collected in the Carolinas, USA. This genus
has been considered as a synonym of Dacrymyces by McNabb (1973), Reid (1974),
and other authors, a point of view that was critically questioned by Oberwinkler (1993).
Some authors, such as Brasfield (1938) and Kennedy (1958), suggested that there is
not sufficient evidence to separate Dacrymyces from Arrhytidia and that further studies
are needed. In the same sense, Martin (1949, p. 85) wrote the following: "The concept
of Arrhytidia here suggested is that, except for its firm gelatinous or waxy texture, it is
similar to Dacrymyces in its earlier stages, beginning as a group of rooted pustules or
discoid fructifications which soon anastomose and eventually form a netted or more
or less continuous film on the substratum." Lowy (1971, 1972, 1975), who undertook
studies on the jelly fungi from the Neotropics, accepted Arrhytidia and described a
new species from Argentina and a new variety from Bolivia. In this study, we consider
Arrhytidia as a valid genus, with the understanding that the taxonomy of species
belonging to this genus needs to be exhaustively studied.
DiStribution: The class Dacrymycetes in Mexico is recorded throughout 24 states (of
32), with Jalisco having the most records, followed by the State of Mexico and Veracruz
(Figs 2–3). The State of Mexico, however, has the most species (13): Arrhytidia
involuta (Schwein.) Coker, Calocera cornea (Batsch) Fr., C. macrospora Brasf.,
C. viscosa (Pers.) Fr., Dacrymyces chrysospermus Berk. & M.A.Curtis, D. dictyosporus
G.W.Martin, D. punctiformis Neuhoff, D. stillatus Nees, Dacryopinax elegans (Berk.
& M.A.Curtis) G.W.Martin, D. lowyi S. Sierra & Cifuentes, D. spathularia (Schwein.)
G.W.Martin, Guepiniopsis buccina (Pers.) L.L.Kenn., and Heterotextus alpinus (Earle)
G.W.Martin. The state’s high species diversity is partly due to two factors: (1) the
proximity of woodlands to the main research centers in the country (Mexico City),
and (2) the State of Mexico has been the main study area in the country for research
specialists on Dacrymycetes, such as Bernard Lowy and Sigfrido Sierra. Hence, more
exemplars from Jalisco, State of Mexico and Veracruz are represented in herbaria.
The vast majority of taxonomic studies that include dacrymycetoid fungi have been
conducted in the central part of Mexico, from the Pacific (Jalisco), to the Atlantic side
(Veracruz), probably due to the high concentration of research facilities in those areas,
followed by southern Mexico, and lastly, northern Mexico with the fewest records for
Dacrymycetes (apart from the states with no records for Dacrymycetes: Aguascalientes,
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Table 2. First geographic records by state and number of records for the 21 species of Dacrymycetes
fungi recorded in Mexico.
Species No. records State First reference
Arrhytidia involuta (Schwein.) Coker2 State of Mexico Lowy 1965
Veracruz Lowy 1971
Calocera coralloides Kobayasi 1 Durango Raymundo et al. 2012
Calocera cornea (Batsch) Fr. 28 Chiapas Lowy 1965
Durango Raymundo et al. 2012
Jalisco Guzmán-Dávalos 2001
Mexico City Lowy 1965
Michoacán Sierra 2000
Morelos Mendiola & Guzmán 1973
Nayarit Sierra & Cifuentes 1993
Quintana Roo Guzmán 1983
State of Mexico Lowy 1965
Tlaxcala Sierra 2000
Veracruz Lowy & Guzmán 1979
Calocera macrospora Brasf. 8 Durango Raymundo et al. 2012
Hidalgo Lowy & Guzmán 1979
Mexico City Lowy & Guzmán 1979
Quintana Roo Guzmán 2003
State of Mexico Lowy & Guzmán 1979
Tlaxcala Sierra 2000
Calocera viscosa (Pers.) Fr. 38 Chiapas Chanona-Gómez et al. 2014
Chihuahua Moreno-Fuentes et al. 1994
Coahuila Mendiola & Guzmán 1973
Durango Raymundo et al. 2012
Jalisco Téllez-Bañuelos et al. 1988
Mexico City Tovar-Velasco & Valenzuela
2006
Michoacán Díaz-Barriga et al. 1988
Morelos Lowy & Guzmán 1979
Oaxaca Welden & Guzmán 1978
State of Mexico Guzmán 1972
Tamaulipas García-Jiménez & Guevara-
Guerrero 2005
Tlaxcala Sierra 2000
Veracruz Welden & Guzmán 1978
Cerinomyces lagerheimii (Pat.)
McNabb 1 Quintana Roo Sierra & Cifuentes 2009
Dacrymyces capitatus Schwein. 7 Durango Raymundo et al. 2012
= D. deliquescens var. ellisii (Coker)
L.L.Kenn. Mexico City Lowy 1965 (as D. ellisii)
= D. ellisii Coker Nuevo León Welden & Lemke 1961 (as
D. ellisii)
= Ditiola nuda Berk. & Broome Oaxaca Welden & Guzmán 1978 (as
D. deliquescens var. ellisii)
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Tabasco Lowy 1965 (as Ditiola
nuda)
Dacrymyces chrysospermus Berk. &
M.A.Curtis 54 Chihuahua Sierra 2000
= D. palmatus Bres. Coahuila Lowy & Guzmán 1979 (as
D. palmatus)
Durango Mendiola & Guzmán 1973
(as D. palmatus)
Guerrero Lowy & Guzmán 1979 (as
D. palmatus)
Hidalgo Lowy & Guzmán 1979 (as
D. palmatus)
Jalisco Guzmán-Dávalos & Fragoza
1995 (as D. palmatus)
Mexico City Mendiola & Guzmán 1973
(as D. palmatus)
Michoacán Sierra & Cifuentes 1993 (as
D. palmatus)
Morelos Lowy 1965 (as D. palmatus)
Nayarit Sierra 2000
Nuevo León Marmolejo & Méndez-
Cortés 2007
Puebla Welden & Lemke 1961 (as
D. palmatus)
Sonora Pérez-Silva & Esqueda-
Valle 1992 (as D. palmatus)
State of Mexico Sierra & Cifuentes 1993 (as
D. palmatus)
Tlaxcala Sierra 2000
Veracruz Lowy & Guzmán 1979 (as
D. palmatus)
Dacrymyces dictyosporus
G.W.Martin 39 Chiapas Lowy 1965
Chihuahua Moreno-Fuentes et al. 1994
Guerrero Sierra & Cifuentes 1993
Jalisco Lowy & Guzmán 1979
Mexico City Lowy & Guzmán 1979
Michoacán Lowy & Guzmán 1979
Morelos Mendiola & Guzmán 1973
Nayarit Sierra & Cifuentes 1993
Nuevo León Lowy & Guzmán 1979
Oaxaca Mendiola & Guzmán 1973
Querétaro García-Jiménez et al. 1998
Sonora Pérez-Silva & Esqueda-
Valle 1992
State of Mexico Lowy 1965
Tlaxcala Sierra 2000
Dacrymyces punctiformis Neuhoff 8 Chiapas Lowy 1965
State of Mexico Lowy 1965
Hidalgo Lowy & Guzmán 1979
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Sonora Pérez-Silva & Esqueda-
Valle 1992
Dacrymyces stillatus Nees 15 Michoacán Díaz-Barriga et al. 1988 (as
D. deliquescens)
= D. deliquescens (Bull.) Duby Morelos Valenzuela et al. 2004 (as
D. deliquescens)
Nuevo León Marmolejo & Méndez-
Cortés 2007
Sonora Pérez-Silva & Esqueda-Valle
1992 (as D. deliquescens)
State of Mexico
Tamaulipas Lowy 1965 (as D. deliques-
cens)
García Jiménez &
Valenzuela 2005 (as D. deli-
quescens)
Dacrymyces variisporus McNabb 1 Mexico City Villarruel-Ordaz &
Cifuentes 2007
Dacryopinax elegans (Berk. &
M.A.Curtis) G.W.Martin 36 Campeche Guzmán 1983
Guerrero Salinas-Salgado et al. 2012
Jalisco Herrera-Fonseca et al. 2002
Michoacán Sierra & Cifuentes 1993
Morelos Lowy & Guzmán 1979
Nayarit Sierra & Cifuentes 1993
Oaxaca Welden & Guzmán 1978
Puebla
Quintana Roo Lowy & Guzmán 1979
Guzmán 1983
State of Mexico Lowy & Guzmán 1979
Tabasco Lowy 1965
Veracruz Lowy 1965
Yucatán Guzmán 1983
Dacryopinax lowyi S. Sierra &
Cifuentes 20 Mexico City Sierra & Cifuentes 2005
Michoacán Sierra & Cifuentes 2005
Morelos Sierra & Cifuentes 2005
State of México Sierra & Cifuentes 2005
Tlaxcala Sierra & Cifuentes 2005
Dacryopinax spathularia (Schwein.)
G.W.Martin 89 Campeche Chio & Guzmán 1982
Chiapas Lowy 1965
Chihuahua Quiñonez-Martínez &
Garza-Ocañas 2015
Guerrero Sierra & Cifuentes 1993
Jalisco Lowy 1965
Mexico City Lowy 1965
Michoacán Díaz-Barriga et al. 1988
Morelos Mendiola & Guzmán 1973
Nayarit Sierra & Cifuentes 1993
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Nuevo León Marmolejo & Méndez-
Cortés 2007
Oaxaca Guzmán 1973
Puebla Lowy 1965
Querétaro Sierra 2000
Quintana Roo Pompa-González et al. 2011
Sonora Esqueda-Valle et al. 1995
State of Mexico Lowy 1965
Tamaulipas García-Jiménez &
Valenzuela 2005
Veracruz Welden & Lemke 1961
Dacryopinax yungensis Lowy 1 Sonora Pérez-Silva & Esqueda-
Valle 1992
Ditiola radicata (Alb. & Schwein.) Fr. 7 Hidalgo Varela & Cifuentes 1979
Morelos Sierra 2000
Quintana Roo Guzmán 1983
Tabasco Lowy 1971
Veracruz Sierra 2000
Guepiniopsis buccina (Pers.)
L.L.Kenn. 10 Durango Lowy 1971
Guerrero Sierra 2000
Hidalgo Sierra & Cifuentes 1993
Mexico City Lowy & Guzmán 1979
Morelos Lowy & Guzmán 1979
State of Mexico Lowy & Guzmán 1979
Veracruz Welden & Guzmán 1978
Heterotextus alpinus
(Earle) G.W.Martin 18 Durango Raymundo et al. 2012 (as
Guepiniopsis alpina)
= Guepiniopsis alpina (Earle) Brasf. Hidalgo Sierra & Cifuentes 1993 (as
Guepiniopsis alpina)
Morelos Sierra 2000 (as Guepiniopsis
alpina)
Puebla Sierra 2000 (as Guepiniopsis
alpina)
Querétaro Sierra 2000 (as Guepiniopsis
alpina)
State of Mexico Sierra & Cifuentes 1993 (as
Guepiniopsis alpina)
Tlaxcala Sierra & Cifuentes 1993(as
Guepiniopsis alpina)
Veracruz Sierra 2000 (as Guepiniopsis
alpina)
Heterotextus luteus (Bres.) McNabb 3 Veracruz Sierra 2000
Heterotextus peziziformis (Berk.)
Lloyd 3 State of Mexico Sierra 2000
Veracruz Sierra 2000
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Baja California, Baja California Sur, Colima, Guanajuato, San Luis Potosí, Sinaloa,
and Zacatecas). Most records for Dacrymycetes are found in Lowy’s publications,
which included 91 records distributed in 15 states. These numbers are followed by 85
records of Guzmán in 16 states and 76 records of Cifuentes in 10 states.
phenology: Although, most species of Dacrymycetes have been collected throughout
the year (a characteristic shared with other groups of fungi in the country, e.g., Chacón
& Guzmán 1995), herbarium records for the species are more common for the summer
months (June, July, August, and September), which correspond to the rainy season
in Mexico (Fig. 4). From the total number of records (389), 199 had associated
phenological data. The two months with the greatest number of records (July and
August) also show the highest species diversity: 82 records and 15 species in July and
44 records and 12 species in August.
Dacrymyces chrysospermus and Dacryopinax spathularia have been sampled
throughout much of the year, from April to December and from March to November,
respectively. The records for Calocera coralloides Kobayasi, C. macrospora, C. viscosa,
Dacrymyces capitatus Schwein., D. dictyosporus, D. punctiformis, Dacryopinax lowyi,
D. yungensis Lowy, and Heterotextus alpinus correspond with the rainy season (July
to September). Ditiola radicata (Alb. & Schwein.) Fr. has been recorded solely in
November. Other species have a more irregular record history, which probably reflects
the uneven sampling of species during the year. For example, Dacrymyces stillatus
is recorded in January, June, July, and August; Dacryopinax elegans in February,
May, July, August, September, and November, and Guepiniopsis buccina in June,
July, September, and December. Calocera coralloides was considered as a "branched
form" of Calocera cornea by McNabb (1965) and considered as "species inquirendae"
Fig. 1. Number of species and records for the eight genera of Dacrymycetes recorded in Mexico.
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because the holotype was destroyed during the Second World War. We consider that
McNabb’s (1965) suggestion is true because only one record was found in Mexico.
The oldest dacrymycetoid record in Mexico corresponds to Dacryopinax spathularia
collected in October 1854 (see Guzmán 1973). Since the study of Sierra in 2000, there
has been more interest in studying Dacrymycetes (see Table 1 for an exhaustive list).
Vegetation typeS: Most species of Dacrymycetes have been reported in pine (Pinus
spp.), fir (Abies religiosa (Kunth) Schltdl. & Cham.), and oak (Quercus spp.) forests,
followed by tropical evergreen, tropical deciduous, and cloud forests. All species were
found growing in fir forests, with the exception of four: Calocera coralloides, collected
only in Pseudotsuga-Populus forests (Kobayasi 1939); Cerinomyces lagerheimii
(Pat.) McNabb, only in tropical evergreen forests (Sierra & Cifuentes 2009); Ditiola
radicata in pine-oak mixed forests and tropical evergreen forests, and Dacryopinax
yungensis only in pine forests (Pérez-Silva & Esqueda-Valle 1992). Although Ditiola
radicata has been described as a species restricted to conifer forests (Reid 1974), one
record (Guzmán without data, ENCB, XAL) indicates that this species is also found
in tropical evergreen forests (Guzmán 1983).
Dacryopinax spathularia is the most widespread species, with records in 16 different
vegetation types, including conifer-oak forests, tropical evergreen forests, tropical
deciduous forests, shrub, coffee plantations, and cloud forests. On the other hand,
Fig. 2. Geographic records of Dacrymycetes in Mexico. Numbers surrounding the map correspond
to geographic coordinates. Dots correspond to Dacrymycetes records.
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Dacrymyces punctiformis, as well as Ditiola radicata, have been found mainly in
conifer forests with a single record in a tropical evergreen forest (Pérez-Moreno &
Villarreal 1988). Dacrymyces punctiformis had been previously cited as exclusively
inhabiting conifer forests (Reid 1974, Shirouzu et al. 2009).
Arrhytidia involuta, Calocera macrospora, Dacrymyces variisporus McNabb,
D. stillatus, and Heterotextus alpinus are recorded exclusively for pine, fir, and oak
forests. Dacrymyces stillatus has been considered a widespread cosmopolitan species
(Reid 1974, Oberwinkler 2014). Calocera cornea, C. viscosa, Dacrymyces capitatus,
D. dictyosporus, Dacryopinax elegans, D. lowyi, D. spathularia, and Guepiniopsis
buccina have been collected in cloud forests and other vegetation types.
The members of the class Dacrymycetes have a characteristic saprobic habit, growing on
angiosperm and gymnosperm wood without a defined pattern. Nevertheless, Shirouzu
et al. (2014) show that dacrymycetoid fungi are more efficient in degrading wood
of Pinus species than wood of other species such as Quercus or Fagus. Most of the
records in Mexico correspond to species growing within conifer forests, supporting
the observations of Gilberston (1980) and Shirouzu et al. (2012) that brown-rotting
fungi are more frequent in conifer forests. Shirouzu et al. (2014) considered that
the preference for pine is the result of a greater degrading capability and a complex
interaction between biotic and abiotic factors. In the case of Calocera, C. cornea
preferentially grows on wood of many widespread angiosperms, whereas C. viscosa
grows mainly on conifer wood in temperate-cold forests (Reid 1974, Shirouzu et al.
2009, Oberwinkler 2014).
Fig. 3. Number of species and herbarium records by state for Dacrymycetes recorded in Mexico.
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Discussion
The study of the taxonomy and species diversity for many groups of fungi in Mexico
remains insufficient. In the case of Dacrymycetes, most inventories usually include
taxa from this group, but relatively few studies focused exclusively on them. In a study
of the ultrastructure of the basidiospores in some Dacrymycetales in Mexico, Sierra
et al. (2013) effectively differentiated among taxa using the presence or absence of
septum pores and the number of layers in the walls of the basidiospores.
According to Kirk et al. (2008) and Shirouzu et al. (2013), 9–10 genera and 110–113
species of Dacrymycetes have been described. These authors do not consider Arrhytidia
and Dacryomitra Tul. & C.Tul. (sensu Oberwinkler 2014) as valid genera. Thus, a more
thorough systematic and phylogenetic study of Arrhytidia is underway by Sierra and
collaborators to validate the inclusion of this taxon within the Dacrymycetes. Eight of
the 11–12 genera, corresponding to 21 species, have been recorded in Mexico (Table
2), which accounts for 18% of the extant diversity of Dacrymycetes.
Compared with Mexico (covering approximately 2 million km²), other regions of the
world have been more extensively studied. For example, Reid (1974) cites 20 species
of Dacrymycetes for the British Islands (approximately 315,000 km2) alone. Because
Mexico has a great variety of habitats in which Dacrymycetes species probably occur,
it is necessary to intensify sampling efforts across the country and to exhaustively
review herbarium specimens to uncover potentially new species. The lack of taxonomic
knowledge in Mexico is not exclusive to Dacrymycetes; this lack is a widespread
problem for many groups of fungi, many of which are solely known from fungal
inventories. For instance, from Psathyrella (Fr.) Quél., a common agaricoid fungus
Fig. 4. Phenology of Dacrymycetes basidiomes in Mexico. The numbers correspond to the records
found.
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378
throughout the world tropics (more than 400 species), only 20 species have been
recorded in Mexico, reflecting the lack of systematic studies for the group (Guzmán
2008). Guzmán emphasized that, in 2008, there were only two monographic studies
for fungi in Mexico [Psilocybe (Fr.) P.Kumm. and Scleroderma Pers.] and highlighted
the lack of specialists as the principal hurdle to the advancement of mycological
knowledge in Mexico. Recent monographic studies include one on Helvella L. by
Landeros & Guzmán-Dávalos (2013), Pluteus Fr. by Rodríguez (2013) Melanoleuca
Pat., by Sánchez-García et al. (2013) and Inonotus P.Karst. by Valenzuela et al. (2013).
The present review shows that the best-represented genera in Mexico, Calocera,
Dacrymyces, and Dacryopinax, correspond to those with the highest species diversity
worldwide. Dacryopinax spathularia is the most common species and exclusively
causes brown rot (Worrall et al. 1997). However, Dacrymyces punctiformis and Ditiola
radicata, two species with a well-known distribution in conifer forests around the
world, are represented in Mexico by a single record in tropical evergreen forests. A
similar case is that of Dacryopinax yungensis, a species reported for a tropical forest in
Bolivia (Lowy 1961) that is recorded for Mexico in a pine forest in the state of Sonora.
Thus, the identity of these species in herbarium records must be carefully reviewed.
The identity of many other records is also doubtful and may need to be revised, as is
the case of Calocera coralloides, a species only registered from one Mexican locality.
In conclusion, the diversity and taxonomic knowledge of Dacrymycetes in Mexico is
poor, in part because many regions in the country have been poorly explored. More
species of Dacrymycetes will undoubtedly be discovered as these regions are more
thoroughly explored. Currently, the Trans-Mexican Volcanic Belt is the best-known
area. For a better understanding of species diversity of this group, taxonomic studies
and fieldwork are necessary in many places, particularly in tropical and subtropical
regions and states with no records (Aguascalientes, Baja California, Baja California
Sur, Colima, Guanajuato, San Luis Potosí, Sinaloa, and Zacatecas). To date, the 21
species recorded in Mexico are mainly distributed throughout conifer forests. Extensive
taxonomic studies are also needed to validate or determine the identity of existing
herbarium specimens.
Acknowledgements
David Espinosa and Alfredo Justo made many suggestions on a draft version of this manuscript.
Othón Alcántara drew the map, Patricia González-Ávila helped us to resolve distributional doubts,
and Ernesto Chanes Rodríguez Ramírez helped with graphs. DGAPA, UNAM, and PAPIIT (project
IV201015) and Consejo Nacional de Ciencia y Tecnología (CONACyT 413867) are thanked for
financial support. The first author acknowledges the Programa de Posgrado en Ciencias Biológicas
for a grant. This study fulfills a requirement for the first author for the degree of Doctor in Sciences
in the Posgrado en Ciencias Biológicas, UNAM.
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Manuscript submitted October 10, 2016; accepted March 10, 2017.
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