ECOGRAPHY 29: 57
/65, 2006

Seasonal community structure of macromycetes in Veracruz, Mexico

P. Munguia, G. Guzman and F. Ramrez-Guillen

Munguia, P., Guzman, G. and Ramrez-Guillen, F. 2006. Seasonal community

structure of macromycetes in Veracruz, Mexico.
/ Ecography 29: 57
/65.
A study of phenological patterns in macromycete communities in Veracruz, Mexico
was carried out in order to understand changes in community structure across regions
of different vegetation types. Previous studies suggest that similarities in community composition occur when there are similarities in certain geographical and
climatological characteristics, however they do not address functional groups or
seasonal changes across regions. Macromycete communities in Veracruz showed similar
species distribution patterns, but individual assemblages changed structure seasonally,
changes that were strongly correlated with rainfall. Interestingly, the number of
functional groups (species performing similar ecological functions) was not determined
by rainfall, but the distribution of species within functional groups was determined by
rainfall. Temperature did not appear to play a role in structuring community diversity
at this regional scale. However, temperature and other environmental factors such as
pH or light may be the mechanism triggering phenological patterns, and influencing the
species pool at localized scales. This work brings new light to fungal community
diversity patterns in a largely unknown group of species.
P. Munguia, (munguia@bio.fsu.edu), Dept of Biological Science, Florida State Univ.,
Tallahassee, FL, 32306-1100, USA.
/ G. Guzman and F. Ramrez-Guillen, Inst. de
Ecologa, Apartado Postal 63, Xalapa, Veracruz, 91000, Mexico.

One of the main goals in ecology is to determine the

mechanisms that affect diversity at different spatial
scales (Brown and Lomolino 1998). Species interactions
such as competition and facilitation can determine both
the abundance and distribution of species at local or
relatively small spatial scales. Alternatively, climatic
factors such as rainfall and temperature can determine
species distribution at broader scales. It has been shown
that a combination of these factors can be responsible
for species diversity (Shmida and Wilson 1985), and their
study can lead to the understanding of species commonness and rarity (Rabinowitz 1981). However, the combination of mechanisms that give rise to species
distributions has been a heated debate (Nee et al. 1991,
Hubbell 2001). The main conflict occurs between the
processes affecting dispersal and niche differentiation
among species, giving rise to different community
patterns across regions (Brown 1984, Collins and Glenn

1991, Scheiner and Rey-Benayas 1997). Here we describe

how phenological patterns and functional group diversity affect the distribution of macromycete species in
Veracruz, Mexico.
Ecological studies on mycological diversity and distribution have traditionally been underrepresented in the
literature (Hawksworth 1991). The ephemeral basidiomata of most macrofungi that present fluctuations in
fructification along with insufficient taxonomic knowledge make ecological studies of these organisms difficult
(Seinfert 1981, Eveling et al. 1990, Arnolds and Jansen
1992); however, studies start to infer ecological patterns
utilizing taxonomic databases (Ponder et al. 2001,
Guzman et al. 2003, Munguia et al. 2003).
When communities are compared across regions,
scientists often assess similarities and differences into
different abiotic and biotic patterns (Watling 1981,
Cornell and Lawton 1992, Zobel 1997, Arita and

Accepted 9 June 2005

Copyright # ECOGRAPHY 2006
ISSN 0906-7590
ECOGRAPHY 29:1 (2006)

57

Rodrguez 2002). Assessing data in this manner is

particularly useful when studying large databases or
when an experimental approach has not been taken. For
example, climatic conditions have been shown to affect
macromycete phenological patterns and may limit species distributions (Widden 1981, Frankland et al. 1996).
Other environmental conditions, such as vegetation may
affect the amount and type of resources available for
fungi (Hering 1966, Watling 1981, Guzman 1994).
Alternatively, even if climatic and environmental patterns are similar among regions, community structure
(species composition and their abundances) may differ,
due to species interactions or dispersal ability. Although
different species may be found in different regions,
similar functional groups may be present, suggesting
that the species identity is not as important for
structuring a particular community as its ecosystem
function (Loreau 2000). However, fruiting phenology
may also fluctuate in regions that undergo seasonal
changes. Therefore temporal changes may also contribute to our understanding of the mechanisms that drive
the abundance and distribution of species.
In general, macromycetes produce fruit bodies according to the season or climatic conditions (Arnolds and
Jansen 1992, Chacon and Guzman 1995). This seasonal
production has been well documented, and it has been
shown that species can differ in their seasonal occurrence
patterns among different regions (Munguia et al. 2003).
Studies at a local scale on hypogenous fungi have shown
that occurrence patterns are different in autumn and
spring seasons (Claridge et al. 2000). However, at a
biogeographic scale seasonal changes in community
composition have not been recorded. Taking into
account seasonal changes in macromycete community
studies is particularly important because most fungi tend
to be identified by their ephemeral fruit body, even
though the mycelia are present year round. If these
seasonal patterns are not considered in studies, then they
may miss many species that are part of the community.
The purpose of this study was to understand macromycete community structure in different regions of the
state of Veracruz in eastern Mexico. We analyzed a
taxonomic database as a hypothesis-generating method
to further ecological studies on macromycetes. This
study addresses several questions in hierarchical fashion
focusing on climatic and vegetative patterns, community
structure, and species identities: 1) Are species distributions different among regions or habitats? 2) Are there
seasonal changes in species fruiting patterns within
regions? 3) Do regions have community structures
different from random assemblages throughout the
year? 4) What is the relationship between functional
groups and community structure across seasons? In this
study we focus on rainfall as the main influential climatic
factor because preliminary analyses have shown that for
58

these particular scales (i.e. regional) temperature presents no effect.

Methods
Description of data
We used data from the taxonomic database of the
Inst. de Ecologa, Mexico (XAL; Guzman et al. 2003).
Eight regions in the state of Veracruz, were chosen
for this study: 1) Veracruz
/Papaloapan, 2) Los Tuxtlas
/
Catemaco, 3) Uxpanapa
/Coatzacoalcos 4) Xalapa
/
Rafael Lucio, 5) Zoncuantla, 6) Coatepec
/Xico, and
7, 8) two distinctive regions within the Cofre de Perote
area (Fig. 1). Each region was separated by at least 15
km (mean /60 km). These regions were also part of an
altitudinal gradient. The regions were grouped into three
main vegetation types: tropical forest from sea level to
900 m (regions 1
/3 above), subtropical cloud forest from
900
/1700 m (regions 4
/6), and two alpine forests
composed primarily of either a) Pinus and Quercus trees
(1800
/2800 m; region 7), or b) Abies trees (2800
/3000
m; region 8). In each type of forest, secondary vegetation
(meadows or shrubs) was also considered where the
original vegetation is under high human influence
(regions 1
/6). Within each region, an average of 25
localities were sampled, except region 5 where five
localities were intensively studied relative to the others.
All eight regions presented similar temperature and
rainfall patterns although the magnitude varied considerably among regions (Fig. 2).
The fungi in the database were collected from 1970 to
2000. Each sample in the database represents an
individual macromycete with one or more fruit bodies,
and the data for such individual includes species name,
date and site of collection, collector, habitat and vegetation type. Macromycete sampling occurred year round,
however, emphasis has been placed during the rainy
season (June
/September) because that is when the
majority of species produce fruit bodies. Therefore,
during the dry season the regions have been sampled
proportional to the wet season. The number of records
for the eight regions comprises 84% of the total number
of records in the XAL database. A total of 5627 records
comprised of 693 macromycete species were analyzed in
this study. These species were mainly comprised of
Ascomycetes (e.g. Pyrenomycetes and Discomycetes)
and Basidiomycetes (e.g. Agaricales, Aphyllophorales,
Gasteromycetes s.l., and Tremellales; where Agaricales
and Aphyllophorales were the most abundant).

Data analysis
The first analysis consisted of comparing fungal distribution across the eight regions (question 1). Previous
ECOGRAPHY 29:1 (2006)

Rainfall (mm)

Fig. 1. The eight regions under

study in the state of Veracruz,
Mexico. Tropical forest: 1)
Veracruz
/Papaloapan, 2) Los
Tuxtlas
/Catemaco, 3)
Uxpanapa
/Coatzacoalcos;
subtropical cloud forest: 4)
Xalapa
/Rafael Lucio, 5)
Zoncuantla, 6) Coatepec
/Xico;
alpine forest: 7) Cofre de Perote
with Abies forest, and 8) Cofre
de Perote with Pinus and
Quercus forest.

500
450
400
350
300
250
200
150
100
50
0

10

11

12

10

11

12

Month

Te mperature (C)

30

25
20
15
10
5
0
1

Month

Fig. 2. Mean monthly rainfall a) and temperature b) for the

eight regions in this study. Thick line represents the mean of the
eight regions9/95% confidence intervals.
ECOGRAPHY 29:1 (2006)

studies have shown that at the regional scale, many

species tend to be sparse, occurring on few localities, and
a small group of species occurs in several local areas
within the region; further, species abundances tend to
have a positive correlation with the number of occupied
sites (Brown 1984, Scheiner and Rey-Benayas 1997).
Here, we wanted to compare species distributions and
see if they varied across regions or vegetation types. We
correlated the number of sites occupied by each species
with its abundance for each region. We also compared
the relationship between fungal site frequency (number
of sites occupied) and local richness (number of species)
among the eight regions. In the case of any differences,
this would suggest that processes at the level of either
region or vegetation could produce differences in the
distribution and local abundance of species.
We were also interested in whether or not fungal
fruiting pattern changed throughout the year (question
2). Fungal fruit bodies vary in the length of time they are
present in the vegetation. Therefore, it is interesting to
note how variation in fruiting phenology can change
apparent species composition (i.e. above ground) at
different seasons within a region. Further, by comparing
community structure during different seasons, we can
infer major changes in above-ground species composi59

60

across seasons (question 4). We divided the mushroom

species into 10 habitat utilization types: corticolous,
ectomycorrhizal, endomycorrhizal, fimicolous, folicolous, fungicolous, terricolous, humicolous, hypogeous,
and lignicolous. A functional group index was created
using Simpsons index of diversity (Magurran 1988)
where functional diversity /Sp2i , where pi is the proportion of species in the ith functional group. Both the
number of functional groups and Simpsons index were
regressed against the bimonthly rainfall for each region.
All statistical analyses were performed using SAS 8.01
(SAS Inst., Cary, NC, USA) and CANOCO (Centre for
Biometry Wageningen, Wageningen, the Netherlands).

Results
Species abundance and distribution
Most of the species analyzed in this study are sparsely
distributed, occurring at only a few sites within a region
(Fig. 3). In fact 63% of the species occurred at only
one site within each region. In all eight regions, there
was a positive correlation between a species abundance
and the number of sites it was present (Kendalls Tau,
mean correlation/0.66, SD /0.22, all p-values B/0.05).
Wright (1991) noted that such a correlation could be due
to sampling artifacts. In order to test for sampling
effects, we regressed the natural logarithm of species
absence frequency (ln(1/p), where p is the percentage of
sites occupied) against the mean number of species
within a region. Under the Poisson null hypothesis,
this relationship should have a slope of /1 and intercept
of 0. For our dataset, the slopes across all regions ranged
from /0.05 to /0.2, and the intercepts ranged from
0.99 to 2.0 (all intercepts statistically different from
zero). Therefore, sampling artifacts do not seem to be
responsible for the positive correlations we observed.
Pairwise correlations were performed among all regions
to assess similarities in the frequency-by-site distributions. The distributions were positively correlated across
all regions (Kendalls Tau, mean correlation /0.84,
SD /0.12, all p-values B/0.05). In other words, all
Proportion of species

tion within a region. We performed a seasonal comparison of community structure using Correspondence
Analysis (CA) with a scaling on intersample distances
(bi-plot scaling) and a down-weight of rare species. The
matrix consisted of 441 species grouped into bimesters
for each region. Because region 4 had more species than
the other regions, we rarefied the species to the mean
abundance levels of these seven remaining regions. We
grouped monthly observations into bimesters in order to
minimize the number of zeros in each sampling unit, and
excluded those species that only occurred once. The first
three axes scores from the CA were then compared using
a MANOVA with either bimonthly periods or vegetation
as treatments. In addition, we performed a second CA
on region 4, Xalapa
/Rafael Lucio, with its full complement of species to illustrate how community structure
can change seasonally within a region.
Fungal community structure at the regional scale was
tested against a null model to determine whether
communities were assembled at random (question 3).
A community matrix with species presence-absence was
created for each bimester for each region. The rows of
each matrix represent the species, while the columns
represent the bimesters within each region. Each matrix
was analyzed separately using the C-score index (Stone
and Roberts 1990, 1992, Gotelli 2000). The C-score
index measures the average number of checkerboard
units between all possible pairs of species. A checkerboard unit is any sub-matrix of the form [0 1, 1 0]:
species A present in region 1 and species B present in
region 2. The larger the resulting C-score value, the less
likely there is a pair-wise species co-occurrence. Each
species occurrence is randomly re-shuffled within each
row of the matrix, and the model ranks the regions
suitability relative to the observed species richness
ranking. Caution should be taken when interpreting
these results because the C-score index may confound
aggregation and segregation in large matrices (Stone and
Roberts 1992). However, in our study, we were not
testing for aggregation or segregation of fungal species,
but departures from random community structure.
Therefore for regions that were not randomly assembled,
the C-score should be significantly smaller or larger than
expected by chance. We performed the C-score analysis
for each bimester with 1000 iterations using Ecosim
(Gotelli and Entsminger 2001). The difference between
the observed and expected C-score values gives the
degree of randomness or non-randomness, if the difference is small or large, respectively. We then tested
the effect of the environment (rainfall and vegetation
type) in structuring regional species assemblages. Each
bimonthly difference in C-score values (observed
/
expected) was regressed against the average rainfall for
the respective time period.
The final analysis consisted of testing the relationship
between functional groups and community structure

1
0.8
0.6
0.4
0.2
0
0-20%

21-40%

41-60%

61-80%

81-100%

Proportion of sites

Fig. 3. Species frequency distribution for the eight regions.

Black bars represent tropical forests, white bars represent cloud
forests, and thatched bars alpine forests. Bars are means for the
regions within each forest type /1 SE.
ECOGRAPHY 29:1 (2006)

regions presented a similar unimodal species frequency

distribution curve, with a median of one site.

Community structure and species identity

Although the eight regions presented similar species
frequency distributions, the species that occurred in
more than eight localities were not the same in more
than two regions (see Table 1 for a list of the most
common species in each of the vegetation types). Only
six species were found to occur in more than eight sites
(i.e. relatively common) in more than one region:
Discomycetes: Helvella macropus ; Aphyllophorales:
Lentinus crinitus, Pycnoporus sanguineus and Schizophyllum commune, Agaricales: Amanita gemmata and A.
rubescens. However, no species were common in more
than two regions.
Community composition, species identity and abundance, varied across vegetation types as shown by the
Correspondence Analysis (Fig. 4a, Table 2). Within each
bimester, community composition differed among alpine
forest and tropical and cloud forest. However, during the
first three bimesters (dry and early wet season), tropical
forests and cloud forests had similar community compositions (Tukeys HSD test on first two axes scores,
p/0.05). In the last three bimesters, late wet season to

the dry season, tropical forests and cloud forests were

statistically different (p B/0.05).
When comparing seasonal community composition
within each vegetation type, only the alpine forest
showed seasonal differences in CA scores (Table 2).
These scores were grouped into early (bimesters 1
/3)
and late (bimesters 4
/6) seasons. However, caution may
be required when interpreting these results, since the
seasonal variation may be occurring at smaller spatial
scales rather than at the vegetation level. Due to low
replication, the analysis was performed on each vegetation using the CA scores for all the regions. Visual
analysis can only be performed on individual regions.
Only region 4 has been extensively sampled, and we
present its seasonal variation in species composition
as an example of regional responses across seasons
(Fig. 4b). A correspondence analysis on region 4 grouping scores by bimester shows a cyclical pattern in
community structure.
In order to test whether or not community structure is
assembled at random for each bimester and if different
vegetation types present different assembly patterns, we
calculated the C-score index in two different ways. First,
we calculated C-scores that corresponded to each
bimester for each vegetation type, providing 18 different
scores (3 vegetations types /6 bimesters). We regressed

Table 1. A list of the most common species in the studied regions.

Regions 1
/3 (tropical)
Cookeina speciosa (Fr.: Fr.)
Dennis
C. tricholoma (Mont.) Kuntze
Cordyceps melolonthae var. rickii
(Lloyd) Mains
Daldinia eschscholzii (Ehrenb.: Fr.)
Rehm
Dictyophora indusiata (Vent. ex Pers.)
Desv.
Earliella scabrosa (Pers.) Gilb. &
Ryvarden
Fomitopsis feei (Fr.) Kreisel
Ganoderma colossum (Fr.) C.F. Baker
Hexagonia papyracea Berk.
Lentinus velutinus Fr.
Lycogalopsis solmsii E. Fisch.
Marasmius cladophyllus Berk.
M. haematocephallus Mont.
Nothopanus eugrammus (Mont.)
Singer
Oudemansiella canarii (Jungh.) Hohn.
Phellinus linteus (Berk. & M.A.
Curtis) Teng
Phillipsia domingesis Berk.
Pycnoporus sanguineus (L.: Fr.)
Murrill
Trichaptum perrottetii (Lev.) Ryvarden
Trogia buccinalis (Mont.) Pat.

ECOGRAPHY 29:1 (2006)

Regions 4
/6 (subtropical)

Amanita bisporigera G.F. Atk.
A. jacksonii Pomerleau
A. virosa Lam.: Secr.

Regions 7
/8 (alpine)

Amanita gemmata (Fr.)
Bertillon
A. rubescens (Pers.: Fr.) Gray
A. muscaria (L.: Fr.) Pers. ex Hook.

A. yema Guzman & Ramrez-Guillen

Aleuria aurantia (Fr.) Fuckel

Boletus ananas (M.A. Curtis) Murrill

Boletus luridus Schaeff.: Fr.

Calostoma cinnabarinum Desv.

Citocybe gibba (Pers.: Fr.) P. Kumm.

Cordyceps gracilis Mont. & Dur.

C. militaris (L. ex St. Amans) Link
Cryptothecia rubrocincta (Ehrenb.) Thor
Daldinia concentrica (Boton: Fr.) Ces. &
De Not.
Entoloma murrai (Berk. & M.A. Curtis)
Sacc.
Gymnopilus lepidotus Hesler
Gyroporus castaneus (Bull.: Fr.) Quel.
Hypholoma subviride (Berk. & M.A.
Curtis) Dennis
Lactarius indigo (Schwein.: Fr.) Fr.

Fomitopsis pinicola (Swartz: Fr.) P. Karst.

Gomphus floccosus (Schwein.) Singer
Gyromitra infula (Schaeff.: Fr.) Quel.
Helvella macropus (Fr.) P. Karst.

Lentinula boryana (Berk. & Mont.)

Pegler
Psilocybe mexicana R. Heim
P. zapotecorum R. Heim emend.
Guzman
Rigidoporus ulmarius (Sow.: Fr.)
Imazeki
Veligaster nitidum (Berk.) Guzman &
Tapia

Hygrophoropsis aurantiaca (Wolfen: Fr.)

Maire
Hygrophorus chrysodon Batsch: Fr.
Hypholoma fasciculare (Huds.: Fr.) P. Kumm.
Inocybe fastigiata (Schaeff.: Fr.) Quel.
Neolentinus suffrutescens (Brot.: Fr.) May &
Wood
Panaeolus semiovatus (Sow.: Fr.) Lundell &
Nann.
Russula cyanoxantha Schaeff.: Schwein.
R. delica Fr.
Suillus brevipes (Peck) Kuntze
Tricholomopsis rutilam (Schaeff.: Fr.) Singer

61

Fig. 4. a) Correspondence
analysis among regions. Each
data point represents a single
bimester in each region,
therefore representing the
seasonal variation within a
given region as well as the
spatial variation among
vegetation types (N/48 data
points). b) Correspondence
analysis of region 4 (Xalapa
/
Rafael Lucio). Each data point
represents the community
structure at a given bimester,
represented by the numbers
(e.g. 1/January
/February).

a
Tropical forest

+1.0

Alpine forest

b
5

Cloud forest

6
4

1
-1.0

the difference in observed and expected C-score indices

against the mean bimonthly rainfall (Fig. 5a). The bestfit line shows a significant concave up relationship
(DF /17, F/5.39, R2 /0.42, p/0.017). At low rainfall
levels community structure is not different from a
random structuring, at intermediate rainfall communities become more structured and at high rainfall levels
communities appear random again. This approach
suggests that fungi found in all vegetation types undergo
similar seasonal changes. However, only cloud forests
presented a significant relationship with respect to
rainfall (DF /5, F/44.09, R2 /0.92, p/0.003). Both
tropical forests and alpine forests presented no relationship (p /0.05).

-1.0

-1.0

It may be argued that this concave relationship was

due to a larger sampling effort during the intermediate
bimesters when rainfall is high. Therefore, we performed
a second C-score analysis that included all regions across
all vegetations for each bimester giving us six different
scores (i.e. one for each bimester). These comparisons
tested whether fungal communities in different regions
presented patterns different from random assemblies for
each bimester. If the difference in the observed and
expected C-score index is low, then the communities
across all regions are not structured. Therefore all
species have the potential to occur in any region.
Alternatively, if the difference is high, then communities
represent structured communities, in which species

Table 2. Correspondence analysis and MANOVA results on CA scores for macromycete diversity on eight regions divided into three
vegetation types. Cumulative variance (VAR) explained by the first three axes and eigenvalues (EIGEN) for the first two axes are
shown. VEG/MANOVA performed on each vegetation type using bimesters as treatment. TIME/MANOVA performed on each
bimester using vegetation as treatment. MANOVAs present the F values for each test. TF/Tropical forest; CF/Cloud forest;
AF/Alpine Forest. Degrees of freedom for MANOVAs are VEG/17, TIME/7. Statistical differences at */ B/0.05, **/ B/0.01,
***/ B/0.001.

VAR
EIGEN
VEGETATION TYPE
VEG
BIMESTER
TIME

62

AXIS 1

AXIS 2

AXIS 3

36.8
0.776
TF
0.87
1
8.89**

46.4
0.537
CF
0.73
3
28.55***

47.8
0.483
AF
9.7***
5
19.83**

2
17.56**

4
32.81***

6
33.74***

ECOGRAPHY 29:1 (2006)

0.15
0.12
0.09
0.06
0.03
0

12

Functional groups

C-SCORE (OBS-EXP)

0.18

100

200
Rainfall (mm)

300

10
8
6
4
2
0

400

100

200

300

400

C-Score (OBS-EXP)

0.060

0.050
0.040
0.030

0.020

0.010
0.000
0

50

100

150
200
Rainfall (mm)

250

300

Fig. 5. a) Differences in C-scores regressed against mean

bimonthly rainfall for each region. b) C-score observed versus
expected differences across all eight regions against the mean
bimonthly rainfall. Numbers in (b) correspond to each bimester
in ascending order (e.g. 1/January
/February).

primarily occur in specific regions. If there was a

sampling effect present, as suggested above, then the
difference in C-score values at intermediate rainfall levels
should be higher than the rest. These communities
should present more structure when compared to communities from different vegetation types. The results of
the C-score difference regressed against rainfall showed a
convex relationship (Fig. 4b; DF/5, F/20.8, r2 /0.93,
p/0.017). At low and high rainfall levels regions are not
assembled at random, however at intermediate rainfall,
regions are not different from random assemblages.

Functional groups
The number of functional groups present in a given
region was highly variable across seasons (Fig. 6a). For
example, in the dry months most of the species present
are lignicolous due to the perennial character of the fruit
body (except Pleurotus djamor and other agarics), while
most saprobe and ectomycorrhizal species fruit in the
wet months. However, the number of functional groups
present in a region was not dependent on rainfall
(Fig. 6a; DF/17, F/0.13, p /0.72). When incorporating the number of species per functional group by way of
the Simpson index, there was a significant negative
relationship with rainfall (Fig. 6b; DF/17, F /5.47,
R2 /25.5, p/0.03). In other words, at high rainfall
levels, most species belonged to one or few functional
groups of the many groups that were present. At low
ECOGRAPHY 29:1 (2006)

Functional group diversity

Rainfall (mm)

0.070

1.2

1
0.8
0.6
0.4
0.2
0
0

100

200

300

400

Rainfall (mm)

Fig. 6. Relationship between fungal functional groups and

rainfall. a) Number of functional groups for each region in
each bimester. The relationship is not significant, solid line is for
illustrative purposes. b) Simpsons index of functional group
diversity (i.e. number of species within each functional group)
regressed against rainfall.

rainfall levels, species were more evenly distributed

across functional groups.

Role of temperature
Regions differ in temperature regimes, alpine forests
presenting the coolest monthly mean temperatures and
tropical forest the warmest (11 and 248C respectively). It
is interesting to note that temperature does not have the
same influence on community structure as rainfall. We
conducted the same regressions between C-scores or
functional groups and temperature and we found no
relationship (DF /17, R2 /0.09 and 0.03 respectively,
p /0.4). This is probably due to the small range in
temperatures (Fig. 2), which tend to fluctuate more at a
microhabitat scale than at the regional scale.

Discussion
The results from this study suggest that fungal community structure is dependent on the vegetation type, and
that seasonal rainfall may influence community similarities among regions. Although rainfall and temperature
patterns are similar across all eight regions, the magnitude of each varies throughout the year. The eight
regions presented similar distributions of common and
rare fungal species, regardless of vegetation type. How63

ever, community structure differed, thus distinguishing

alpine from cloud from tropical forests. What was most
interesting was that seasonal changes in rainfall influenced how structured the eight communities were at any
given point in the year. Changes in species assemblages
did not influence the number of functional groups
present. Therefore, vegetation type was the most important component in structuring macromycete communities. As a result of this association between fungi and
plants, fungal species identity appeared to be more
important than their function.
The fungi in this study appeared to be niche-based
(Scheiner and Rey-Benayas 1997): most species occurred
within one or few habitats in a region (Fig. 3), regardless
of the vegetation type. Furthermore, there did not seem
to be a single, common distribution pattern among the
most common species, different species were found in
different regions. Because there were no distribution
differences across regions, the species pool present at a
particular region seemed to depend on vegetation
(Fig. 4). However, rainfall was an influential factor
structuring diversity across all regions regardless of
vegetation type. Niche-based fungal guilds appeared to
strongly depend on the seasonality driven by rainfall
(Fig. 5).
During intermediate rainfall levels fungal species seem
to fruit at random, while at low rainfall levels only those
species with perennial fruit bodies persisted and formed
a non-random assemblage. At high rainfall levels, species
may fruit in sequence, depending on their niche requirements (Munguia et al. 2003). This fruiting pattern
created non-random assemblages across regions of
different vegetation types. However, within each region
communities only appeared as random assemblages,
which could be due to fluctuations in microhabitat
conditions.
These seasonal changes in community structure are
important to consider when assessing macromycete
diversity. Previous work on macromycete communities
has focused on relationship between diversity and
environmental variables (e.g. vegetation, temperature
and rainfall) or the relationship between phenology
and environmental variables (Watling 1981, Bills et al.
1986, Luoma et al. 1991). When using fruit bodies to
measure diversity, it is important to consider fungal
phenology because the observed species pool at one
moment in time may actually underestimate the actual
number of species present, yet unseen. At large spatial
scales, seasonal fluctuations of fruit body diversity
would reflect such phenological patterns. Fruit body
production depends on several environmental factors
including rainfall, temperature, light and soil pH (Hunt
and Trappe 1987, Luoma et al. 1991). In our study
rainfall appeared more important than temperature at
regional scales. The role of temperature may be to trigger
fruiting, or it may be more important at smaller spatial
64

scales (e.g. within 100 m) or different latitudes. Soil pH

poses a more difficult factor to incorporate in our
studies at such large-scale.
As we have demonstrated, macromycete distribution is
determined by vegetation and environmental variables.
Studies such as this one show phenological patterns,
which could be a subset of the total mycological
assemblage. However, microhabitat factors such as light
and temperature may determine how species identities
and phenologies will vary across different vegetation
types or even regions. At the microhabitat scale, we
suggest competition for resources as well as timing of
reproduction will create different observed species pools
while community structure remains the same across
regions. It is important to note however that at these
large scales, random processes associated with dispersal
(Hubbell 2001) may still explain the abundance and
distribution of species.
Acknowledgements
/ We are grateful to L. Guzman-Davalos,
A. Hollebone, B. Inouye, and T. Miller for discussions and
revisions of the manuscript. Guzman and Ramrez-Guillen
thank Etelvina Gandara, Mara Eugenia Ramrez, Manuel
Hernandez and Juan Lara for their help in maintaining the
database and herbarium work. Financial support for the
database was through CONABIO (Project E006).

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Subject Editor: Francisco Pugnaire.

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