Research Article

Tropical Conservation Science

October-December 2016: 1–17
Distributional patterns of the Order ! The Author(s) 2016
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DOI: 10.1177/1940082916667140
in Mexico trc.sagepub.com

Patricia Astrid González-Ávila1, Isolda Luna-Vega1,

Ricardo Garcı́a-Sandoval1, and Raúl Contreras-Medina2

Abstract
The distribution and endemicity patterns of Gomphales in Mexico are analyzed here for the first time. Richness and cor-
rected endemism were obtained from a dataset of 3,483 records for 97 species, using a cell-grid system of one degree per
side. The central region of Mexico (Tlaxcala, Veracruz, Hidalgo and Estado de México states), which includes most of the
Trans-Mexican Volcanic Belt and Sierra Madre Oriental biogeographic provinces, had the highest richness values, but Mexican
areas in the Pacific Coast (Jalisco and Michoacán states) and southeastern Mexico (Oaxaca and Chiapas states) had the most
distinctive composition, as measured by the corrected endemism index. Two main distributional patterns were recovered: a)
montane: at elevations above 1000 m on coniferous, pine-oak, oak and cloud forests, typified by the presence of species of
Ramaria and Clavariadelphus, b) lowlands: at elevations below 1000 m mainly in evergreen, rainforest and deciduous tropical
forests characterized by the presence of four tropical species of Lentaria, two tropical species of Gomphus endemic to
Mexico, and four tropical species of Phaeoclavulina. The eight species of Gomphales endemic to Mexico have very restricted
distribution, mostly in non-protected areas, and are not considered under special protection programs. The present con-
tribution delineates general patterns of distribution for the Gomphales, and documents its diversity and endemism in Mexico.

Keywords
Agaricomycetes, Basidiomycota, Phallomycetidae, neotropic, endemism

catalogue of the Mexican fungi commissioned by the

Introduction
Mexico is one of the most biologically diverse countries in
the world (Flores-Villela & Gerez, 1994; Martı́nez-Meyer,
Sosa-Escalante, & Álvarez Noguera, 2014; Mittermeier &
Goettsch de Mittermeier, 1992; Ramamoorthy, Bye, Lot,
& Fa, 1993), due to factors such as its geographical pos-
ition, geological history, climate and orography. It is also
a transition zone between the Nearctic and Neotropical
biogeographic regions (Halffter, 1987). The geographic
distribution of the elements comprising Mexican biota
is the result of vicariance, local dispersion and extinction
events as well as Pleistocene climatic changes and in situ
speciation processes (Salinas-Moreno et al., 2004).
A relevant, but often neglected, part of this biodiver-
sity is fungi. Mexican fungal diversity has been estimated
at between seven thousand (Guzmán, 1998a, 1998b) and
almost eleven thousand species (Aguirre-Acosta, Ulloa,
Aguilar, Cifuentes, & Valenzuela, 2014). A preliminary
Comisión Nacional para el Conocimiento y Uso de la
Biodiversidad (CONABIO, www.conabio.gob.mx) has
documented more than 4,700 species for the main
fungal groups (Garcı́a-Sandoval et al. unpublished
data). So far, a comprehensive catalogue of Mexican
fungal diversity is not available, but some estimates and
1
Laboratorio de Biogeografı́a y Sistemática, Departamento de Biologı́a
Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México
(UNAM)
2
Laboratorio de Botánica, Escuela de Ciencias, Universidad Autónoma
‘Benito Juárez’ de Oaxaca (UABJO)
Received 10 December 2015; Revised 4 July 2016; Accepted 6 July 2016
Corresponding Author:
Isolda Luna-Vega, Laboratorio de Biogeografı́a y Sistemática, Departamento
de Biologı́a Evolutiva, Facultad de Ciencias, Coyoacán, Mexico 04510,
Mexico.
Email: luna.isolda@gmail.com

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2 Tropical Conservation Science
preliminary catalogues exist for specific groups (i.e.,
González-Ávila, Luna-Vega, Villegas-Rı́os, Saade, &
Cifuentes, 2013; González-Ávila, Torres-Miranda,
Luna-Vega, & Villegas-Rı́os, 2013; Herrera-Campos
et al., 2014).
Some studies on fungal distribution have been con-
ducted in Mexico, although drawing on incomplete
data. For example, Cifuentes et al. (2004) and
Gónzalez-Ávila, Luna-Vega, et al. (2013) analyzed the
distributional data of 214 species in the Sierra Madre
Oriental in Mexico applying a parsimony analysis of
endemicity or PAE (Rosen, 1988). Their study did not
find a significant correlation between fungal and plant
distributions, instead, they reported a more significant
relation between fungal composition and physiographic
features. Similar findings relating fungal distributions to
non biological parameters were reported by Tedersoo
et al. (2014).
The Gomphales is one of the fungal orders with high
relevance in Mexico, where edible mycorrhizical and
saprophytic species are reported frequently (González-
Ávila, Luna-Vega, et al., 2013). Most of these species
have great traditional and economic importance as food-
stuff in some states of Mexico. The members of this
group can be found in several vegetation types, including
tropical, subtropical and temperate forests, from sea level
to 3900 m (González-Ávila, Luna-Vega, et al., 2013;
González-Ávila, Torres-Miranda, et al., 2013). The
three families considered in the order, following Kirk,
Cannon, Minter, and Stalpers (2008), are present in
Mexico, and 10 out of 18 genera have been reported in
the country (González-Ávila, Luna-Vega, et al., 2013;
González-Ávila, Torres-Miranda, et al., 2013). Ramaria
is the genus with most species (55) recorded in Mexico,
followed by Phaeoclavulina (16), Clavariadelphus (8),
Gomphus (5), Gautieria, Gloeocantharellus, Lentaria and
Turbinellus (2 each), and Beenakia and Kavinia with only
one species each. Until 2013, a total of 92 species were
reported from Mexico, which corresponds to 27% of the
total known diversity of the group in the world
(González-Ávila, Luna-Vega, et al., 2013).
In Mexico, the species of Gomphales are mainly found
in coniferous forests, where Abies, Pseudotsuga, Picea
and Pinus are the dominant genera (Estrada-Torres,
1994). However, some species have also been reported
in tropical forests, and even some endemic species are
distributed in these types of vegetation (Villegas-Rios,
Cifuentes, Estrada-Torres, & Kong-Luz, 2010). From
an anthropocentric point of view, these fungi are relevant
because several ethnic groups use them as part of their
diet and some taxa are ectomycorrhizal species related
mainly to coniferous genera (Aguilar, & Villegas-Rı́os,
2010; González-Ávila, Luna-Vega, et al., 2013).
In some countries the inclusion of plant and animal
(and fungi) species in protection programs or on
Red Lists usually requires information about population
structure and dynamics (IUCN, www.iucn.org). In the
specific case of fungi, and due to the difficulties inherent
in obtaining information regarding, for example, popula-
tion size or generation length, their conservation mostly
relies on evidence of their distribution (Dahlberg, &
Mueller, 2011; Molina, Horton, Trappe, & Marcot,
2011).
In the case of Mexico, Norma Oficial Mexicana 059
(NOM-059, www.profepa.gob.mx, SEMARNAT, 2010)
represents the Mexican Red List of endangered species.
This list incorporates 49 species of fungi, all of them
lacking proper justification for their status because, in
Mexico, there is no formal protocol for the risk evalu-
ation of fungal species. This situation produces uncer-
tainty about the real status of the species on the list,
and results in a high possibility of these species not
being considered as really at risk. In Mexico, Red Lists
are used as complementary criteria for the delimitation of
protected areas (Urquiza-Haas, Tobón, & Koleff, 2011).
It is therefore necessary to have appropriate protocols
and criteria for the inclusion of fungal species on those
lists, and to have reliable information on distribution,
richness and endemicity of endangered species.
The present study intends to: a) provide a first formal
analysis on the distribution of species of Gomphales in
Mexico, identifying areas with the highest species richness
and endemism, and b) provide the first evaluation of the
conservation status for the group in Mexico, identifying
species with restricted distribution and their relation to
protected areas.
Methods
Distributional data
Distributional records of the Gomphales species in
Mexico were obtained from four main sources: a)
Specialized literature: papers published in peer-reviewed
journals with an emphasis on taxonomic studies; b)
Public databases of government environmental agencies
(i.e. www.conabio.gob.mx), or databases associated with
mycological collections, both private and public; c) Field
work: a series of field trips were conducted to collect
specimens in areas identified as poorly sampled; d)
Mycological collections: distributional records were
obtained directly from specimens deposited in the main
Mexican herbaria.
The main bibliographic sources for distributional data
were: González-Ávila, Luna-Vega, et al. (2013);
González-Ávila, Torres-Miranda, et al. (2013); Estrada-
Torres (1994); Villegas-Rı́os et al. (2010); Aguilar y
Villegas-Rı́os and Villegas-Rı́os (2010); Castillo, Garcı́a,
and San Martı́n (1979); León and Guzmán (1980);
Martı́nez, Pérez-Silva, and Aguirre-Acosta (1983);
González-Ávila et al.
Guzmán (1983); Zarco (1986); Dı́az-Barriga, Fefer, and
Valenzuela (1988); Villegas-Rı́os and Cifuentes (1988);
Cifuentes et al. (1990); Montoya (1997); Nava and
Valenzuela (1997); Montoya, Estrada-Torres, Kong-
Luz, and Juárez (2001); Montoya, Hernández-
Totomoch, Estrada-Torres, Kong-Luz, and Caballero
(2003); Montoya, Kong-Luz, Estrada-Torres, Cifuentes,
and Caballero (2004); Herrera, Guzmán-Dávalos, and
Rodrı́guez (2002); Valenzuela-Garza, Pérez-Ramı́rez,
and Morales-Torres (2004); Garibay-Orijel (2006);
González-Ávila (2006, 2010); Landeros Castillo,
Guzmán, and Cifuentes (2006); Chanona-Gómez,
Andrade-Gallegos, Castellanos-Albores, and Sánchez
(2007); Villarruel-Ordaz and Cifuentes (2007); González-
Ávila, Villegas-Rı́os, and Estrada-Torres (2013).
The following mycological collections were consulted:
The Mycology Herbarium at the Universidad Autónoma
of the state of México, CHIP, ENCB, EBUM, FCME,
FEZA, The Mycology Herbarium at the Universidad
Autónoma of Morelos, IBUG, IZTA, MEXU, TLXM,
UJAD, CIAD, ITCV and XAL (Index Herbariorum:
http://sweetgum.nybg.org/).
Fieldwork was carried out in the Mexican states of
Chiapas, Distrito Federal, Durango, México, Guerrero,
Hidalgo, Michoacán, Oaxaca and Tabasco. The herbar-
ium specimens from mycological collections and our field
samples were identified at species level, with the exception
of several specimens of the genus Ramaria, which were
only identified at subgenus level, mainly due to a lack of
morphological data, particularly coloring, which is neces-
sary for correct taxonomic identification.
Figure 1. Grid-cells in which at least one taxon of Gomphales was recorded in Mexico.
3
Distribution analyses on richness and endemism
Richness and endemism analyses were conducted on
three datasets: a) Core dataset (CDS) of 987 records for
56 species. Records in this dataset correspond to speci-
mens identified at species level or specimens recognized as
putative independent species, either because they may
correspond to species not formally described or because
specimens were poorly documented in the field (i.e. lack-
ing color notes, which was the case for most of the spe-
cimens from Ramaria). B) Extended core dataset
(ExCDS): the core dataset was extended to include rec-
ords of Ramaria identified at the subgenus level following
Marr and Stuntz (1973). Records were coded as absence
or presence of the subgenera in the grid-cell. C) Genera
dataset (GDT): this set includes records from the previ-
ous datasets, but the basic units for analysis were genera
following Giachini and Castellano (2011).
The analyses based on the datasets were conducted
with a grid of one degree per side dividing the Mexican
territory (Figure 1). We decide to use this grid size
because it facilitated data manipulation, reducing sam-
pling artifact effects (such as mapping errors and
grid-cells where sampling did not take place in sparsely
inhabited areas), and because it has been reported that
smaller sizes may disaggregate the patterns of richness
and endemism (Crisp et al., 2001; Laffan and Crisp,
2003; Linder, 2001).
Species endemism was calculated on the grid-cell ana-
lysis following the proposal of Crisp et al. (2001) and
Linder (2001), using weighted endemism and the
4 Tropical Conservation Science
corrected weighed endemism indexes. In the case of the
first index, the value of each species was obtained and
weighted according to the inverse of its distribution, so
that if a taxon is only present in one grid-cell, it is given a
value of 1, whereas if a taxon is registered in two grid-
cells its value is 0.5 and so on. To obtain the value of each
grid-cell, the values of all the species in the grid-cell were
added together. Therefore, grid-cells containing many
species showing restricted distribution are expected to
have higher values than those grid-cells with many
widely-distributed species. To obtain the corrected
weighted endemism index, the weighted endemism value
of each grid-cell was divided by the total number of spe-
cies in that grid-cell. This operation ‘‘corrects’’ species
richness (Crisp et al., 2001) so that grid-cells containing
relatively few species, but showing restricted distribution,
should register higher values than those grid-cells with
high richness, but containing relatively few species with
restricted distribution. Although the total number of spe-
cies in each grid-cell was also taken into consideration for
corrected weighted endemism, unlike weighted endem-
ism, this should not be dependent on richness, and a
low correlation between the two variables was expected
(Crisp et al., 2001; Linder, 2001; Santa Anna del Conde,
Contreras-Medina, & Luna-Vega, 2009).
Grid-cell analysis, maps and index calculations were
performed using Biodiverse software v. 0.99 (Laffan,
Lubarsky, & Rosauer, 2010). In order to evaluate the
level of correlation between weighted endemism and cor-
rected weighted endemism values with species richness,
we estimated the r2 using a simple linear model, and a
simple Pearson coefficient. Even if this method is not
appropriate for this kind of data, we decided to include
the results to enable better comparability with previous
studies (Crisp et al., 2001; Linder, 2001; Santa Anna del
Conde, Contreras-Medina, & Luna-Vega, 2009). Results
from the richness analysis were clustered in Biodiverse
0.99 (Laffan et al., 2010) using Sorensen’s index, and
results were plotted on a map according to the groups
recovered.
Additionally, an analysis of the number of species and
genera present was done using biogeographic provinces
(Arriaga, Aguilar, Espinosa, & Jiménez, 1997) and
Mexican states as geographic units (Figure 2). We
decided to use states as units because in Mexico, as well
as in other countries, conservation policies are generally
conceptualized taking political boundaries into consider-
ation, rather than natural criteria (Contreras-Medina &
Luna-Vega, 2007; Dávila-Aranda, Lira-Saade, & Valdés-
Reyna, 2004).
Species and genera were categorized as Nearctic,
Neotropical, Holarctic, Pantropical, endemic and
Cosmopolitan depending on their known distribution.
An additional map was generated using a digital layer
of the Protected Natural Areas of Mexico (CONABIO,
www.conabio.gob.mx). Distribution records for species
restricted to Mexico or first described in Mexico
were plotted on the map in order to visually inspect
their representation in the National network of
Protected Natural Areas.
Results
Distributional datasets and general patterns
The database assembled included a total of 3,483 records,
comprising 10 genera and 97 species of the order
Gomphales (Figure S1), distributed in 81 grid-cells
(Figure 1). The core dataset (CDS) included 56 species
and it was the only one considered for endemism analysis,
because the units of comparison are similar, which
enabled us to avoid potential distortions resulting from
the subjectivity of taxonomic ranking (Bertrand, Pleijel,
& Rouse, 2006), or from considering non monophyletic
taxa (Szumik & Goloboff, 2015). Distributional patterns
for the genera dataset (Figure S1) showed a high concen-
tration of species in central and southeastern Mexico
(with the exception of Lentaria), broadly in agreement
with the Mexican mountain ranges (mostly in the center
of the country, where the Trans-Mexican Volcanic Belt is
located, Figure 2b).
Global patterns of distribution
The elevational patterns described in the analysis support
the previously reported relationship between distribution
of ectomycorrhizal species and elevation (Bahram,
Polme, Kõljalg, Zarre, & Tedersoo, 2012). Our results
also support a relationship with vegetation types, but
this may also be related to the distributional patterns of
the associated plants (Miyamoto, Nakano, Hattori, &
Nara, 2014). The present study represents
the first attempt to categorize the country with reference
to fungal distributions, particularly with Gomphales
species.
Patterns recovered by the CWE index are less sensitive
to distortions produced by richness patterns or bias in the
sampling effort (Figure 3). Notwithstanding, they remain
sensitive to under-sampled taxa and operational ende-
mics. To our knowledge this is the first time that the
use of the CWE has been documented for correcting sam-
pling bias.
Some Mexican states with coniferous and oak forests,
such as Aguascalientes, San Luis Potosı́ and Sinaloa,
have few or no records of Gomphales. This is probably
due to poor sampling rather than a real absence of
Gomphales in those forests. Unfortunately, there are no
institutions in these states where mycology research has
been developed, and this has led to gaps in knowledge
about the fungal diversity of the country.
González-Ávila et al. 5

Figure 2. (a) The 32 Mexican states; b. The 19 biogeographic provinces of Mexico according to Arriaga et al. (1997). Abbreviations are:
AGS ¼ Aguascalientes, BC ¼ Baja California, BCS ¼ Baja California Sur, CAMP ¼ Campeche, CHI ¼ Chiapas, CHIH ¼ Chihuahua,
COA ¼ Coahuila, COL ¼ Colima, DF ¼ Distrito Federal, DUR ¼ Durango, GTO ¼ Guanajuato, GRO ¼ Guerrero, HGO ¼ Hidalgo,
JAL ¼ Jalisco, MEX ¼ México, MICH ¼ Michoacán, MOR ¼ Morelos, NAY ¼ Nayarit, NL ¼ Nuevo León, OAX ¼ Oaxaca, PUE ¼ Puebla,
QR ¼ Quintana Roo, QRO ¼ Querétaro, SLP ¼ San Luis Potosı́, SIN ¼ Sinaloa, SON ¼ Sonora, TAB ¼ Tabasco, TAMP ¼ Tamaulipas,
TLA ¼Tlaxcala, VER ¼Veracruz, YUC ¼ Yucatán, ZAC ¼ Zacatecas; (b) the 19 biogeographic provinces of Mexico according to Arriaga
et al. (1997). Abbreviations are: apn ¼ Altiplano Norte, aps ¼ Altiplano Sur, bal ¼ Depresión del Balsas (Balsas Depression), bc ¼ Baja
California, clf ¼ California, cab ¼ Del Cabo, chi ¼ Los Altos de Chiapas (Chiapas Highlands), gm ¼ Golfo de México (Gulf of Mexico),
nus ¼ Soconusco, oax ¼ Oaxaca, pac ¼ Costa del Pacifico (Pacific coast), ptn ¼ Petén, sme ¼ Sierra Madre Oriental, smo ¼ Sierra Madre
Occidental, sms ¼ Sierra Madre del Sur, son ¼ Sonorense, tam ¼ Tamaulipeca, vol ¼ Eje Volcánico (Trans-Mexican Volcanic Belt),
yuc ¼ Yucatán.
6 Tropical Conservation Science
Figure 3. Map of Mexico and dendrogram showing two main clusters of Mexican Gomphales, a montane and a lowland group.
The differential sampling problem results in areas with
apparently different richness patterns, regardless of their
similarity in vegetation types. This situation needs to be
considered before arriving to final conclusions concern-
ing diversity patterns in the country.
We detected at least six general distribution patterns
in the Mexican species of Gomphales. These were:
Holarctic (38 species), Nearctic (31 species),
Cosmopolitan (9 species), Pantropical (6 species),
Neotropical (5 species), and endemic to Mexico (8 spe-
cies). These general patterns of distribution confirm
their association with temperate vegetation, mainly
with northern affinities (Table S1).
The Sorensen’s index-based dendogram supported a
clear distinction between montane and lowland regions
(Figure 3). These patterns are partially related to the
vegetation types and the elevation with which that par-
ticular fungal species is associated.
The montane group included grid-cells related to con-
iferous, pine-oak, oak and cloud forests, on elevations
above 1000 m, and located mainly in the Trans-Mexican
Volcanic Belt and the Sierra Madre Oriental. Prevalent
species in this group are those of Ramaria and
Clavariadelphus.
The lowland group included grid-cells related to trop-
ical vegetation types, such as evergreen and deciduous
forests (but not necessarily the same across the grid-
cells). The corresponding grid-cells are distinguished
by the presence of four tropical species of Lentaria
(L. surculus and three additional species that have not
been described, which are endemic to Mexico), two spe-
cies of Gomphus endemic to Mexico (G. albidocarneus and
G. calakmulensis), and four species of Phaeoclavulina
(P. articulotela, P. gigantea, P. insignis and P. zippelli).
Two non-contiguous grid-cells were distinctly dissimi-
lar to the rest (AAA and FFF). These grid-cells were
located in lowlands (one on the Pacific coast and the
other on the Gulf of Mexico). The first included parts
of the states Michoacán and Guerrero, and the second
parts of the states of Veracruz, Oaxaca and Puebla. These
grid-cells do not share fungal species and do not seem to
be associated with any recognizable pattern.
Distributional analyses
Species level core dataset (CDS). The richest grid-cell was the
TT (19 species), located in the central region of the country,
comprising most of the state of Tlaxcala, including La
Malinche National Park (Figure 4a). The second richest
grid-cell was the KK (16 species), which partially includes
two states and two National Protected Areas (Table 1). The
following grid-cells with the highest values are located in the
central and southeastern parts of the country, and are asso-
ciated with Protected Natural Areas (Figures 4a and 5).
Patterns recovered from the weighted endemism index
(WE) were similar to richness patterns, as reported pre-
viously (Crisp et al., 2001). The grid-cell with the highest
value was TT (10.705), followed by grid-cells SS (6.7165)
and KK (6.4596). These three grid-cells had the greatest
richness as well (Figure 4a,b).
On the other hand, the corrected weighted endemism
index (CWE) showed a different pattern compared to
either richness or weighted endemism (Figure 4c). Grid-
cells showing the highest values are AAA (1.0) located in
the south of Michoacán, Y (0.69) located in Jalisco and
Aguascalientes, and FFF (0.75) covering parts of
Veracruz, Oaxaca and Puebla.
Grid-cell AAA includes only one record corresponding
to a species of Phaeoclavulina which has not been
described. The grid-cell FFF includes the only record of
Beenakia fricta in Mexico and one of only two records of
P. roellinii. The grid-cell Y (on the border of
Aguascalientes and Jalisco) includes records of two spe-
cies of Phaeoclavulina and one of Clavariadelphus trunca-
tus, neither of which are formaly described (i.e. the
species has not been described and properly published
according with the code of nomenclature). The latter spe-
cies has a wide distribution.
González-Ávila et al.
Figure 4. Highest richness, weighted endemism and corrected
weighted endemism values of grid-cells based on the distribution of
species of Gomphales.
Richness and WE showed a very strong correlation
(r2 ¼ 0.9215, p-value: <2.2e-16), while richness had a
low correlation with CWE (r2 ¼ 0.0483, p-value: 0.1174).
Correlations on these indexes in the Gomphales datasets
(Figure S2) are similar to those reported previously for
other taxa in Mexico (Santa Anna del Conde et al., 2009).
When states and biogeographical provinces were used
as comparison units, richness patterns were congruent
with those observed for the CDS. The central and south-
eastern parts of the country had the highest values, and the
7
northern states had the lowest values (Tables 2 and 3),
probably because of a bias in the sampling rather than a
real absence of Gomphales in those areas.
Analysis of the extended dataset (ExCDS). Richness and
endemism patterns recovered from the analysis of the
ExCDS were highly congruent with results from CDS
(results not shown). This could be because the information
from the records of subgenera of Ramaria may be outnum-
bered by the records of the species level dataset. Ramaria
represents the most species-rich genus of Gomphales in the
country (58 species).
We assembled a database of 2,775 records, many of
which were identified only to the subgenus level due to
missing information, because fresh specimens were poorly
documented in the field. The most diverse zones in terms
of number of subgenera, species and/or subspecies of this
genus were grid-cells SS and NNN (Figure S3), located in
the center of the country, corresponding with montane
localities in elevations from 1,000 to 3,900 m, in conifer-
ous, oak and cloud forests (Table 1). Records for
Ramaria subgenera have a richness pattern similar to
other species of Gomphales, with the highest values con-
centrated in the center of the country. Edible and mycor-
rhizal species (Table S2 and Table S3).
Analysis of the genera dataset (GDT). The grid-cell containing
the highest number of genera was UU, containing seven out of
ten of the genera recorded for Mexico (Figure S4). This grid-
cell included Cofre de Perote National Park, on the border of
Veracruz and Puebla (Table 1). The second richest cell was
the TT with six genera, which covers most of the state of
Tlaxcala, including most of La Malinche National Park.
The following richest grid-cells had a pattern of distribution
congruent with that observed in the core dataset, with the
highest values in the center and in some areas of southeastern
Mexico, and a strong association with protected areas.
A similar pattern was recovered with states and bio-
geographic provinces as geographic units of comparison.
The provinces of the Trans-Mexican Volcanic Belt and
the southern part of the Sierra Madre Oriental (Figure 2)
had the highest values of richness of genera (Tables 2
and 3). The regions with fewer records are located in
the northern part of Mexico, most likely because of a
poor sampling effort rather than a real absence of
Gomphales in the area.
Endemic species and their conservation status
The eight species of Gomphales described for Mexico
were located in the center and southeastern part of the
country (Figure 5), consistent with the overall pattern of
richness previously described (Figure 4a). One species
was originally described from La Malinche National
Park (Ramaria bonii), and the protologue is the only
8 Tropical Conservation Science

Table 1. Number of species of Gomphales reported from some of the Mexican Protected Natural Areas (PNAs).

Number
Grid-cell Protected Natural Area Category genera/species

TT Cofre de Perote National Park National Park 7/11

UU – – 6/11
SS Iztaccı́huatl-Popocatépetl and La Malinche National Park 5/21
KK El Chico National Park National Park 5/19
Barranca de Metzitlán Biosphere Reserve Biosphere Reserve
RR Flora and fauna Protection Area of the Flora and Fauna Protection area 5/15
Nevado de Toluca, Desierto de los National Park
Leones National Park, el Tepozteco and
Lagunas de Zempoala National Parks
QQ Mariposa Monarca Biosphere Reserve Biosphere reserve 5/15
AAA Chichinautzin Biological corridor 5/9
VVV Lagunas de Montebello National Park 3/9
DDD – – 3/3

Figure 5. Distribution of endemic species of Gomphales and their relationships with Mexican ANPs.

available record. Four of the species have been recorded
more than once in Protected Natural Areas, but four spe-
cies have one or no records in protected areas (R. persi-
cina, R. radicans, R. suaveolens have no records and
Clavariadelphus fasciculatus have one record).
Sierra Madre Oriental (Figure 2b). This may be partially
the result of the extensive sampling effort in these regions
over several decades (Aguirre-Acosta et al., 2014). In gen-
eral, this pattern of richness follows that reported for
other fungal groups in Mexico (Aguirre-Acosta et al.,
2014).

Discussion
Distributional data and general patterns of richness
The central and southeastern parts of Mexico have the
highest values of richness in the country (Figure 4a), cor-
responding to the Trans-Mexican Volcanic Belt and the
Overall distributional patterns
Most of the species (69 species) of Gomphales known to
be found in Mexico are concentrated in two global pat-
terns of distribution, Holarctic and Nearctic, representing
71% of the species. These two general patterns of
González-Ávila et al. 9

Table 2. Number of genera and species of Gomphales in the Table 3. Number of species of Gomphales in the Mexican states.
Mexican biogeographic provinces.
States Genera Species
Number of Number of
Biogeographic province genera species Aguascalientes 0 0
Baja California Norte 2 3
Altiplano Norte 1 1 Baja California Sur 1 1
Altiplano Sur 5 17 Campeche 3 8
Baja California 0 0 Coahuila 4 5
California 2 3 Colima 1 1
Costa del Pacı́fico (Pacific Coast) 5 9 Chiapas 6 15
Del Cabo 1 1 Chihuahua 3 5
Depresión del Balsas 4 9 Distrito Federal 4 11
(Balsas Depression)
Durango 3 7
Eje Volcánico (Trans-Mexican 7 27
Estado de México 6 19
Volcanic Belt)
Guanajuato 3 5
Golfo de México (Gulf of Mexico) 6 14
Guerrero 4 9
Los Altos de Chiapas 5 7
(Chiapas Highlands) Hidalgo 5 17
Oaxaca 5 7 Jalisco 5 17
Petén 4 15 Michoacán 5 15
Sierra Madre Occidental 3 8 Morelos 5 5
Sierra Madre Oriental 7 21 Nayarit 1 2
Sierra Madre del Sur 5 9 Nuevo León 2 4
Soconusco 0 0 Oaxaca 4 6
Sonorense 0 0 Puebla 5 9
Tamaulipeca 1 1 Querétaro 5 7
Yucatán 1 1 Quintana Roo 4 10
San Luis Potosı́ 0 0
Sinaloa 0 0
Sonora 1 1
distribution confirm their association with temperate Tabasco 3 4
vegetation types such as fir, pine and oak forests. Tamaulipas 4 6
Halffter (1987) noted that the Nearctic and Holarctic pat- Tlaxcala 7 21
terns in insects are related to the mountainous systems of
Veracruz 8 20
Mexico and northern Central America, which have also
Yucatán 1 1
been observed in species of Gomphales. In the case of the
Trans-Mexican Volcanic Belt, this mountain chain has Zacatecas 2 3
been considered a diversification center for animals
(Halffter, 1987) and plants (Styles, 1993); if this situation
is also confirmed for the Gomphales, many more endemic locations. Our results support a closer relationship
species are still to be discovered. between elevation and fungal composition. We were
The altitudinal zones recovered with Sorensen’s index also able to recover a clearer pattern of distribution,
(Figure 3) are congruent with patterns previously probably because our analysis was conducted on a
reported for ectomycorrhizal fungi (Jarvis, Woodward, larger geographical scale.
& Taylor, 2015), where distribution is closely related to Finally, the presence of Beenakia fricta in Mexico
elevation, more than to other parameters, and the great- could be indicative of a cryptic speciation phenomenon
est richness is found above 1000 m.a.s.l. (Miyamoto et al., in this taxon. This species has been reported in Africa,
2014). Europe and Mexico (Borgarino, Moreau, & Richard,
Cifuentes et al. (2004) analyzed the distribution of 220 2005; Nuñez & Ryvarden, 1994). Such a wide distribution
species of macrofungi of the Sierra Madre Oriental is usually associated with lineages composed of multiple
mountain chain through a PAE (Rosen, 1988). These phylogenetic species and low morphological differenti-
authors found a closer relationship between species dis- ation (known as cryptic species). Hawksworth (2012)
tribution and elevation, rather than with vegetation type; argued that this phenomenon is more frequent in taxa
unfortunately, they could not recover a clear pattern of with tropical distributions.
10
Richness and endemicity in the core dataset (CDS)
Grid-cells with the highest richness values (TT, KK and
SS) included Protected Natural Areas (Table 1, Figure 5),
which have traditionally been studied by mycologists
(Aguirre-Acosta et al., 2014). Among the Protected
Natural Areas of Mexico, the Natural Parks and
Biosphere Reserves have been extensively sampled due
to the interest in having a comprehensive inventory of
the species present there. This may have led to a bias in
the richness data, resulting in a distorted pattern of high
richness in areas where there has been extensive sampling
and low richness in poorly sampled areas. These factors
need to be taken into account when considering the rich-
ness patterns recorded for Mexican Gomphales.
Tlaxcala had the most species (21) of Gomphales, des-
pite being the smallest state in surface area in the country.
The states with the next largest numbers of species of
Gomphales recorded were Veracruz (20 species) and
Estado de Mexico (19 species) (see supplementary mater-
ial). These data partially follow the pattern of being the
states where most sampling occurred, as reported by
Aguirre-Acosta et al. (2014).
The results showed that the central and southeastern
Mexican states have the highest values for richness. These
values may reflect the sampling efforts made in certain
states for macroscopic fungi in particular, as mentioned
above. These results may change in the future and are
congruent with patterns of other biological groups of
Mexican biota. Other studies (Flores-Villela & Gerez,
1994; Mittermeier et al., 1992; Ramamoorthy et al.,
1993) have found that the states of Oaxaca, Chiapas
and Veracruz have the greatest biodiversity in the country
and have recently been recognized as megadiverse
Mexican states (Luna-Vega, Espinosa, Rivas, &
Contreras-Medina, 2013).
The biogeographical provinces showing the greatest
diversity of species and genera are the Trans-Mexican
Volcanic Belt and the southern portion of the Sierra
Madre Oriental (Figure 2b). The Trans-Mexican
Volcanic Belt is the richest province for Gomphales. This
result is not surprising, since the Trans-Mexican Volcanic
Belt is one of the most biodiverse zones, where extensive
sampling has occurred, as is corroborated by other groups
of organisms such as Boletaceae fungi (Garcı́a Jiménez J &
Garza Ocañas, 2001), Ternstroemiaceae (Luna-Vega,
Alcántara-Ayala, & Contreras-Medina, 2004) and other
groups of Mexican plants (Contreras-Medina & Luna-
Vega, 2007; Styles, 1993; Villaseñor, 2004; Luna-Vega,
Alcántara-Ayala, Contreras-Medina, & Vargas, 2006), as
well as animals (Fa & Morales, 1993; Garcı́a-Trejo &
Navarro, 2004; Navarro-Siguenza, Lira-Noriega,
Peterson, Oliveras de Ita, & Gordillo-Martı́nez, 2007;
Ochoa-Ochoa & Flores-Villela, 2006).
Biogeographic provinces with no available records are
Baja California, Sonorense and Soconusco. The Baja
Tropical Conservation Science
California peninsula deserves a special mention because
there are some records available in its distal portions
(California and Del Cabo provinces) which are associated
with coniferous forests (Styles, 1993). Most likely the low
richness values reported for this area are due to poor
sampling rather than a real absence of Gomphales in
the area. This may be especially true for regions where
there is a high richness of pine, fir and oak species.
However, in regions such as the Baja California desert,
the lack of proper biotic conditions may be the reason of
the absence of records for Gomphales.
Corrected weighted endemism (CWE) showed a differ-
ent pattern to richness. The main objective for calculating
this index was to avoid the potential distortion produced
by richness values, allowing us to easily recognize areas
where there was a unique or distinctive composition (that
may constitute potential areas of endemism) and that
may be difficult to recognize with other indices. Non cor-
rected endemism index may be very sensitive to an under-
sampling of common species (Crisp et al., 2001), and that
is another reason because of the use of CWE. This index
also allows us to avoid, up to certain level, the distortion
resulting from sampling biases, even when the index may
be sensitive to the under-sampling of common species
(Crisp et al., 2001).
The richness values in Mexican Gomphales showed a
strong correlation with the corrected endemism index and
a low correlation with the CWE. Correlation values
between richness and CWE for Mexican Gomphales
were similar to previous studies with plants, i.e. Crisp
et al. (2001) and Santa Anna del Conde et al. (2009)
and support the use of the CWE as an index relatively
unaffected by richness patterns.
The grid-cell with the highest CWE value was AAA
(1.0), located on the Pacific Coast, on the border between
the states of Michoacán and Guerrero. This grid-cell
includes only one Phaeoclavulina species that has not
been formally described and which is endemic to
Mexico. It inhabits coniferous forest at an elevation of
2,400 m, and was collected from only one locality.
Because this grid-cell has only one record of a species
collected from this locality, the CWE value for the grid-
cell is 1, indicating that the species is restricted to that
single grid-cell (Giachini & Castellano, 2011; González-
Ávila et al., 2013).
This CWE index produces misleadingly high values
when only uncommon species are collected (Crisp et al.,
2001). In our study, high values most likely represent an
absence of data, rather than the relative abundance of
characteristic species. This fact has been reported previ-
ously by Santa Anna del Conde et al. (2009), suggesting
that high values CWE need to be carefully examined to
give accurate results.
The grid-cell FFF (CWE ¼ 0.75), located on the Gulf
Coast, includes two records: the only record of Beenakia
González-Ávila et al.
fricta in Mexico and Phaeoclavulina roellinii, also distrib-
uted in Chiapas (grid-cell WWW). The FFF grid-cell has
a very high value due to the presence of B. fricta, even
when it is a ‘‘false’’ endemic species. B. fricta has been
reported from two other continents (Africa and Europe;
Nuñez & Ryvarden, 1994; Borgarino, Moreau, &
Richard, 2005). The same occurs with P. roellinii
restricted to two Mexican grid-cells, which is also not
endemic to Mexico (it is also distributed in North
America and Europe, according to Petersen (1981)).
The FFF grid-cell contains two records of non-endemic
but highly restricted species in our dataset, producing an
unusually high value. This behavior for the index has
been reported previously (Dávila-Aranda et al., 2004),
but there has been no further discussion.
On the other hand, the grid-cell Y (CWE ¼ 0.68)
includes the only records of two non formally described
species of Phaeoclavulina (potentially endemic to
Mexico), and one record of the widely distributed
Clavaridelphus truncatus. The high value of this grid-cell
is the result of the existence of two restricted and poten-
tially endemic species.
The high CWE values of grid-cells AAA, FFF and Y
are the result of different factors. AAA and Y grid-cells
include endemic species and their values reflect, to some
extent, the aim of the index for recognizing the ‘‘unique-
ness’’ of the grid-cell, but the FFF grid-cell includes what
we call ‘‘operational endemics’’, which are species widely
distributed outside the area of study, but very restricted
in the dataset under analysis.
The presence of ‘‘operational endemics’’ in our distri-
butional data reflects the absence or scarceness of geo-
graphic records or taxonomic studies based on species of
Gomphales in Mexico. In the future, with advances in
Mexican mycological studies, some of the species we
have recorded as false endemics or ‘‘operational ende-
mics’’ (because they are also distributed in other coun-
tries, whereas in Mexico they have only been found in one
grid-cell) are likely to be recorded in other areas or states
of Mexico.
In most of the contributions related to the taxonomy
and biogeography of fungi, the species concept is mor-
phological. However, some studies using microanatomy
or molecular data have shown that some fungi taxa based
on morphology may represent different cryptic species
(Giachini, 2004). In this sense, future studies should
modify the taxonomic status of some of the taxa con-
sidered herein.
Richness and endemicity in the genera
dataset (GDS)
When genera were used as basic units for comparison, the
richness patterns were highly congruent with the previous
ones. All the genera (with the exception of Lentaria and
11
Beenakia as mentioned before) were distributed in the
central and southeastern parts of Mexico, mainly in the
Trans-Mexican Volcanic Belt and in the southern part of
the Sierra Madre Oriental (Figure 2b). Most of the spe-
cies were recorded at elevations up to 3900 m, corres-
ponding with montane areas. These elevations may
partially contribute to the pattern recovered when apply-
ing Sorensen’s index to the richness data (see above,
Figure 3).
Species included in genera as Ramaria, Clavariadelphus
and Gautieria are distributed at the highest elevations
(1000 m - 3900 m), while Lentaria and Beenakia have the
lowest range (10 m-1860 m). Overall, the only pattern that
can be recognized at genera rank indicates that the high-
est richness is associated with higher elevations.
Additionally, there may be some uncertainty if genera
are used as units of comparison, because of the potential
distortions that may result if taxa recognized and for-
mally described under different criteria are compared
(Bertrand et al., 2006), or if non-monophyletic taxa are
considered (Szumik & Goloboff, 2015), which is the case
with Ramaria and Clavaridelphus.
Species with restricted distribution
At least eight species of Gomphales are endemic to
Mexico (Figure 5). Most of the species are distributed
in the Trans-Mexican Volcanic Belt. Three species of
Gomphus have been described from tropical regions in
the states of Campeche, Veracruz and Chiapas, at eleva-
tions from 300 to 1500 m.a.s.l. (Figure 5). It is likely that
more field work sampling at low elevations will discover
more endemic species.
The percentage of endemic species is around 8% (of a
total of 97 species of Gomphales in Mexico). This value is
twice that reported for lichens, according to Herrera
Campos et al. (2014). Building on these authors’ findings,
we can suggest that if the proportion of endemic species is
similar, the number of species of Gomphales in Mexico
could reach 200 species, 59% of the total number of spe-
cies reported by Kirk et al. (2008), or even higher if more
endemic species are discovered.
Implications for conservation
Special protection and conservation programs usually
rely on data from Red Lists. For fungi, those lists are
often based on less information than is available for
plants or animals. Field data for population dynamics
or size is difficult to obtain for most fungal species, but
distributional records are less problematic. Because of
this, conservation policies and programs for fungi usually
rely on distributional records only (Dahlberg & Mueller,
2011). Usually, species with very few records are con-
sidered to have a highly restricted distribution area and
12
to be more susceptible to extinction. For this reason they
are more likely to be included in special protection
programs.
Our paper provides the basic data required to inform
conservation policies for the Mexican Gomphales. These
species meet most of the requirements to be protected and
included on Red Lists and special conservation programs
(Figure 5). There have been very few reports of Ramaria
and Gomphus species in Mexico, and most of those are
from the protologue. A clear example of this is the
Clavariadelphus fasciculatus, a species restricted almost
exclusively to Central Mexico, and which mainly occurs
outside protected areas.
Biogeographical studies, such as the present one, are
especially relevant because, rather than merely generating
data, they provide insights which can inform biology con-
servation policies and programs (Hawksworth, 2012;
Merckx et al., 2014). They also promote a recognition
of centers of richness and endemism (Contreras-Medina
& Luna-Vega, 2007; Crisp et al., 2001).
Another example of unprotected species are those lin-
eages of Gomphales potentially endemic to Mexico. The
eight species described from Mexico are known only in
Mexico, and so far they have not been included in any
protection program (i.e. International Red Lists or
Mexican Norm NOM-059).
Overall, most of the species of Gomphales reported to
have been found in Mexico are distributed in areas with-
out any protection or conservation status. The species
distributed in Protected Natural Areas are present in
just 10% of the total area under protection in the coun-
try, mainly in Central Mexico. Izta-Popo and La
Malinche National Parks, located in the Trans-Mexican
Volcanic Belt, together harbor 19 species.
The apparent high number of species of Gomphales in
National Parks and Biosphere Reserves can be explained
by the extensive sampling that has been carried out in these
categories of Protected Natural Areas. Most of the
Mexican Protected Natural Areas are National Parks,
but the Biosphere Reserves have the biggest area under
protection in Mexico (Flores-Villela & Gerez, 1994).
Some initiatives for conservation of fungal species in
Europe take into account the management and protec-
tion of the habitat (i.e. the forest) as part of a compre-
hensive approach to conservation. The careful
preparation of Red Lists and special programs for ende-
mic species with restricted distribution represents an
essential element of any successful conservation program
(Dahlberg, Genney, & Heilmann-Clausen, 2010;
Dahlberg & Mueller, 2011; Heilmann-Clausen &
Vesterholt, 2008; Molina et al., 2011; Moore, Nauta,
Evans, & Rotheroe, 2008; Senn-Irlet, Heilmann-
Clausen, Genney, & Danhlberg, 2007; Venter et al.,
2014). A comprehensive approach which includes mul-
tiple data sources may be most effective for protecting
Tropical Conservation Science
Mexican Gomphales. Combining the current program
of Protected Areas with a properly elaborated list of spe-
cies at risk may be the most effective approach. Our con-
tribution represents a step in that direction of
conservation biology.
Mexico harbors almost 27% of the known diversity in
the order Gomphales, and probably could reach up to
59% of the known species, if the northern and tropical
regions of the country were more widely explored. The
presence of several potentially endemic taxa has been
documented in temperate areas of Mexico, for example,
in the genus Ramaria subgenus Laeticolora (see Estrada-
Torres, 1994), and the same situation occurs in the tropics
where three species of Gomphus have been described in
recent years (Giachini, 2004).
This region is the most populated and urbanized in the
country and, because of the reduced amount of well con-
served forest, a low richness of fungal species can be
expected. It is likely that the high richness reported is
related to the extensive sampling effort over several dec-
ades. For this reason there is an urgent need to implement
conservation strategies for species known in localities
outside protected areas, such as R. persicina and R. sua-
veolens, as well as those growing in protected natural
areas such as Popo-Izta and La Malinche National Parks.
Acknowledgements
To the people in charge of the collections mentioned in the text for
their courtesy during the review of their specimens; to Adriana
Lomelı́ (UNAM, Mexico) for providing valuable technical assist-
ance; to the Posgrado en Ciencias Biológicas at the Universidad
Nacional Autónoma de México; to Consejo Nacional de Ciencia y
Tecnologı́a (CONACYT), Mexico (the National Science and
Technology Council) for the grant (number 207211) awarded to
the first author. The financial resources for field trips and herbaria
visits came from PAPIIT projects IV201015 and IN215914.
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