Plant Ecology (2005) 180:117–134
Springer 2005

DOI 10.1007/s11258-005-3026-9
-1

Environmental correlates of tree and seedling–sapling distributions

in a Mexican tropical dry forest

Yalma Luisa Vargas-Rodriguez1,2,*, J. Antonio Vázquez-Garcı́a3 and G. Bruce Williamson1

1
Department of Biological Sciences, 107 Life Sciences Building, Louisiana State University, Baton Rouge, LA
70803, USA; 2Current address: Carlos Fuero 543, Colonia Universitaria, Guadalajara, 44840, Jalisco,
Me´xico; 3Centro Universitario de Ciencias Biológicas y Agropecuarias, Departamento de Botánica y Zoo-
logı´a, Universidad de Guadalajara, Km 15 carretera Guadalajara-Nogales, Las Agujas, Zapopan, 45110,
Jalisco, Me´xico; *Author for correspondence (e-mail: yvarga1@lsu.edu)

Received 28 June 2004; accepted in revised form 28 February 2005

Key words: Bray and Curtis ordination, Disturbance, Importance value, Regeneration, Specialization, Tree
diversity, Tropical deciduous forest

Abstract

Bray and Curtis ordination was used to explore which environmental variables explained importance values
and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra
de Manantlán, Jalisco, Mexico. The diameters of trees ‡2.5 cm DBH and the presence and height of
seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data
and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis
ordination. The matrices were based on density, frequency, and dominance of adult trees as well as
seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope,
canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling
species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees
and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1
and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling pres-
ence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting
little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be
important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization
along different soil microsites could promote species coexistence in this forest, while heterogeneity in light
conditions may instead determine differences in growth and, thus, importance value of trees. We
hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely
more important at early stages in tree life histories than in later life history.

Introduction distributed across the Pacific lowlands on the

western side of the country, covering most of
In Mexico, 60% of the tropical vegetation is Jalisco state and confined to small patches in
tropical dry forest (TDF). Mexican TDF is widely eastern Mexico (Rzedowski 1978; Trejo and Dirzo
118
2000). The forest is characterized by marked sea-
sonality in rain fall and its occurrence on moderate
steep slopes and rocky outcrops.
Tropical dry forests in western Mexico, includ-
ing those in the studied area, are typically decid-
uous, with short stature trees and high density of
small size trees (Trejo 1998). Trees ca. 4 m tall
constitute 65%, those with heights between 4 and
8 m are 31% and only 3% are 8–12 m height,
some exceptional individuals reach 15 m. Tree
mean density is 3610 (800) individuals/ha and ba-
sal area is ca. 56.8 m2/ha. Trees with diameter
‡10 cm represent only 20% of individuals and less
than 5% are ‡30 cm, thus, the majority of trees
have diameters £ 2.5 cm (Rzedowski 1978; Trejo
1998). Trees have extended crowns with bright and
peeling barks, as well as compound leaves. Most of
the tree species (ca. 85%) are deciduous and few
(ca. 15%) are evergreen, such as Ziziphus mexicana
Rose and Prosopis laevigata (Willd.) M.C. Johnst.
In addition, columnar and Opuntioideae cacti are
common in this community (Rzedowski 1978;
Trejo 1998).
TDF, throughout Mexico, is severely affected by
livestock, slash and burn agricultural practices,
forest fire, and selective logging. In 1990, only 27%
of the original cover of tropical dry forest re-
mained intact and 1.4% continues to be lost
annually as a result of deforestation (Trejo and
Dirzo 2000). Forest fires and livestock are also
common in Sierra de Manantlán Biosphere Re-
serve (SMBR). Fire in the area has been mainly
associated with agricultural burning, occurring in
the dry season (December–May). Livestock activ-
ity occurs during the rainy season (June–October).
However, forest protection is enforced in core
zones through various mechanisms negotiated
with agrarian communities (68% of reserve’s area)
and private landowners (32% of reserve’s area). In
addition, there are management goals in the buffer
zone directed to implement sustainable practices in
forestry, agriculture, livestock, and other natural
resource management activities (INE 2000).
TDF in Mexico is characterized by high a and b
diversity (Balvanera et al. 2002; Trejo and Dirzo
2002). While species diversity has been shown to
be positively correlated with increasing precipita-
tion across wet and dry forest in Puerto Rico
(Murphy and Lugo 1986), a comprehensive
analysis of Mexican TDF does not show such a
relationship (Trejo and Dirzo 2002). Instead, it has
been suggested that Mexican TDF diversity is
positively related to potential evapotranspiration
(Trejo and Dirzo 2002).
In addition, local site factors such as the
presence and size of canopy gaps, soil properties,
anthropogenic disturbance, and total annual
precipitation may have important effects on TDF
tree species richness, composition, abundance,
and structure (Gonzalez and Zak 1996; Gentry
1988; Oliveira-Filho et al. 1998; Gillespie et al.
2000; Segura et al. 2003). For instance, variation
in species diversity in the Neotropics could be
explained by total annual precipitation (Gentry
1982, 1988). Associations of species with those
factors and specialization to particular micro-
habitats are hypothesized to contribute to species
diversity (Connell 1978; Gentry 1982; Denslow
1987; Welden et al. 1991; Clark et al. 1993).
Different tree species are best suited to different
habitats, which may lead to habitat specializa-
tion. Differential resource utilization might ex-
press itself as microhabitat specialization
between the species or as differences in geo-
graphical distribution. On the other hand, the
availability of different resources may be sepa-
rate in time and may become available during
different seasons. For example, canopy gaps
provide different soil, light, and moisture condi-
tions than the forest understory and could be
exploited by species with specialized competitive
abilities (Denslow 1987; Oliveira-Filho et al.
1998). Differences in species’ growth and survival
may also occur in the absence of gaps along
smaller light gradients, such as between 0.2 and
6.5% available diffuse light (Lieberman et al.
1989; Montgomery and Chazdon 2002). In
addition, natural disturbances can cause spatial
and temporal variability in the availability of
resources leading to habitat differentiation.
Consequently, variation in growth and survival of
tree species under differing conditions of resource
availability (for example micro-topography or
light) could result in habitat specialization (Kobe
1999; Pearson et al. 2003).
Previous studies have shown that microhabitat
specialization with regard to topography and soil
characteristics affect the distribution of several
plant groups, including tropical trees, melasto-
mate shrubs, herbs, pteridophytes, and palms
(Kahn and De Castro 1985; Liberman et al.
1985; Poulsen and Baslev 1991; Basnet 1992;

Toumisto and Roukolainen 1993; Clark et al.
1998; Svenning 1999). For instance, distribution
and abundance of some deciduous tree families
(e.g., Leguminosae), are related to ecological
gradients such as edaphic conditions and shade
tolerance (Givnish 1999). Cation exchange level
affects the distribution of 51 tree species in
Ghana (Swaine 1996). Soil organic matter, soil
nutrients, texture, and moisture are strongly
correlated with main axes of a PCA ordination
of tropical trees in a Bornean tropical rain forest
(Webb and Peart 2000). In addition, adult trees
and seedlings had different strengths of associa-
tion with those variables (Webb and Peart 2000).
Other species can be specialized along moisture
gradients, reflected by the greater numbers of
species that persist on protected side-slope areas
due to their higher dry season moisture levels
(Hubbell 1995).
Even though previous studies used a quantita-
tive multivariate approach to analyze how tree
species relate to environmental variables, only a
few studies have considered the relationship be-
tween the regeneration of tree species in seasonal
tropical dry forest and environmental conditions
(Lieberman and Li 1992; Oliveira-Filho et al.
1998), and none have explored the possible con-
tribution of environmental variables at different
stages in the life cycle of TDF trees.
Here we report the results of a study of tree
forest composition and regeneration along a short
altitudinal gradient in a tropical dry forest at La
Escondida-La Cabaña, Sierra de Manantlán Bio-
sphere Reserve, in the Ayuquila watershed, in
western Mexico, to generate hypotheses about
tropical trees and regeneration patterns along soil
and canopy gaps gradients. We addressed the
following question: What environmental variables
could explain major community gradients for both
trees and seedlings and saplings? To answer this
question, we employed a multivariate gradient
analysis, Bray and Curtis ordination technique
(also known as sociological ordination) (Curtis
and McIntosh 1951; Beals 1984; McCune and
Grace 2002), which produces pure community
gradients that can be correlated with measured
environmental variables. This approach has pro-
vided many insights into the nature, organization
and dynamics of ecological communities (Whit-
taker 1956, 1960; Terborgh 1973; Ludwig and
Reynolds 1988).
119
Methods
Study area
La Escondida-La Cabaña gradient is located in the
northern boundary of the SMBR, roughly 50 km
from the Pacific Ocean, in Jalisco state, Western
Mexico. The study area lies between El Aguacate
(municipio El Grullo) and Zenzontla (municipio
Tuxcacuesco), northwest of ejido Zenzontla, along
the Ayuquila-Armerı́a river, within the tributary
watershed La Pasión-Cerro Blanco. The eleva-
tional gradient extends from Arroyo La Escondida
at, 880 m to the top of Cerro la Cabaña at, 1090 m
a.s.l. (1942¢ N, 10407¢ W) (Figure 1).
La Escondida-La Cabaña occupies the steep
foothills of the northern Sierra de Manantlán, in
the Sierra Madre del Sur, an area of rough
topography, within a hilly and dissected land-
scape. Relief ranges from 10 to 66% slope incli-
nation, along La Escondida and La Cabaña hills.
Microclimate at La Escondida-La Cabaña is
moist, warm, and highly seasonal with a wet sea-
son from June to October and a dry season from
December to May (Martı́nez et al. 1991). Mean
annual rainfall is 900 mm (range 600–1000 mm)
(Martı́nez et al. 1991). Mean annual temperature is
22 C (range 22–28 C), free of frost. The tem-
perature lapse rate is 4 C per 1000 m a.s.l.
(Martı́nez et al. 1991). TDF at La Escondida-La
Cabaña corresponds to the Lower Montane Sub-
humid Tropical Dry Forests of Holdridge (1967)
and occupies 12,700 ha (9%) of the SMBR
(Cuevas-G. et al. 1998).
Tertiary volcanic rocks are prevalent in the
study area (Cruz 1989). Acidic intrusive rocks
(granites) were found on sites 1, 2, 3, 4, 8 and 9,
while intermediate extrusive rocks (rhyolites,
andesites, and trachytes) prevailed on sites 5, 6,
and 7. Land form is convex with boulders and
rocky outcrops.
Shallow, well-drained lime soils (Regosol eut-
rico) were common in sites 5, 6 and 7. Some lime
secondary soils such as Feozem háplico and some
Litosols occured. Litosols (with rocks greater than
7.5 cm in diameter) were frequent at sites 1, 2, 3, 4,
8 and 9. Fluvial sandy soils (Fluvisol eutrico) were
found on river banks and at the foothills of La
Escondida (CETENAL 1975, 1976).
Floristic checklists and descriptions of tropical
dry forest species in the SMBR (Vázquez-Garcı́a
120
Figure 1. Location of La Escondida-La Cabaña at Sierra de Manantlán, Jalisco, Mexico and sampling sites (rectangles).
et al. 1995; Cuevas-G. et al. 1998) contain over
1060 species (38% of the total Sierra de Mana-
ntlán vascular flora). In these checklists, 110
families and 531 genera have been recorded from
tropical dry forest of SMBR. The vegetation is
largely deciduous for up to 8 months, depending
on the length of the dry season. A few species, such
as Ziziphus mexicana and Prosopis laevigata are
evergreen.
Field sampling
The least disturbed forested areas in the Ayuquila
watershed were selected through visual interpre-
tation of topographic maps and field reconnais-
sance. We sampled nine 0.1 ha sites, each one
consisting of ten 2 · 50 m transects, using a
stratified random sampling design along elevation
and exposure to represent variability of the forest.
 121

The total area sampled was 0.9 ha (9 sites · 10

pH 1:2

Tree, shrub and herb gaps quantities represent the averages per site. Total counts of disturbance events and the average of rocks and stones per site are also shown. Soil nutrients
units are ppm. Physiographical unit 1 corresponds to lower slope, 2 to middle slope, and 3 to upper slope. Topography (microrelief shape) is considered in two categories, 1 regular,
6.76
6.76
5.89
6.28
6.28

6.37
6.41
6.64
transects/site · 0.01 ha/transect). Sites were spaced

6.6
at 70 m elevational intervals (Figure 1). This type
%Organic

of sampling allows comparability with the

matter

250 10.15
250 9.13
250 9.47
250 6.43
250 8.12
180 9.12
120 5.07

250 9.81
60 8.12
extensive Gentry’s forest data set (Gentry 1982;
Phillips and Miller 2002). Four sites (1–4) had
southeast exposure, starting near the Ayuquila
NO3 Mg K

river, from foot to top of La Escondida Mountain,

50
50
25
25
50
50
50
50
25
two sites (8–9) were located at the shoulder of the
same mountain, on its eastern slope, three others
12
12
12
12
12

12

25
6

6
(5–7) were located on the west-facing slopes of a
12
12
12
12
12
25
12
25
25
P

neighboring mountain, La Cabaña. The two

Ion exchange capacity

mountains spaced by a small ravine. Slopes

inclination ranged from a 10 to 60% across the
nine sites and all sites had a landform of concave
Exposure Elevation gaps gaps gaps Disturbance Slope Physiography Rocks Stones Topography meq/100 g

shape. At each site, plant and environmental

variables were measured. Diameters of all trees
53.21
51.74
39.08
35.96
35.04
37.98
37.61
33.76
31.93

‡2.5 cm diameter at breast height (DBH, at 1.30 m

height) were measured. Heights of all seedlings
(individuals <130 cm height) and saplings (DBH
<2.5 cm and ‡130 cm height) were recorded in
each of the nine sites (Olvera et al. 1996). Sampling
2
1
2
1
2
2
1
1
2

was conducted from June to October of 1997

(when leaves were present).
4
3
4
3
4
3
2
4
2

Species’ identifications were determined by the

Table 1. Geographic, edaphic, topographic and disturbance variables used in ordinations.

authors except for difficult taxa that were sent to

5
3
4
3
5
2
2
2
2

specialists. Nomenclature generally follows Váz-

quez-Garcı́a et al. (1995). Herbarium specimens
were deposited at IBUG herbaria of the University
of Guadalajara (Holmgren et al. 1990; Holmgren
and Holmgren 1993).
1
2
2
3
1
2
2
1
2

without undulations and 2 corresponds to irregular, with undulations.

Quantitative and qualitative environmental data

65
62
66
58
48
46
24
10
50

were recorded for each transect (Table 1). Eleva-

tion was taken with an altimeter and slope was
measured with a clinometer for each transect.
Exposure was measured with a compass. Topog-
20
20
13
10
21
29
33
37
22

raphy, physiographical unit, and the amount of

Tree Shrub Herb

rocks and stones were registered in each transect.

62
66
66
50
26

10
13
7

Gap presence was determined by the line intersect

method (Battles et al. 1996). Presence of tree,
34
33
20
25
39
57
59
63
61

shrub, and herb gaps were recorded at 5-m inter-

vals along each of the 0.01 ha transects, totaling 10
17
28
34
23
23

11
28
42
9

gap measurements per guild per transect. The

numbers of tree, shrub, and herb gaps were sum-
1020
1090

1060

1000
880
950

920
990

930

marized per site. For gap measurements, we con-

sidered as tree strata all woody plants >2.5 m tall
with their crowns completely or partially exposed
to light; shrub strata all woody plants <2.5 m tall;
SE
SE
SE
SE
W
W
W
E
E

and herb strata all woody and non woody plants

SITE1
SITE2
SITE3
SITE4
SITE5
SITE6
SITE7
SITE8
SITE9

<50 cm tall. While conducting vegetation sam-

pling, two soil samples, 0–30 cm in depth, were
122
taken from random locations at each site. This soil
depth is aimed to obtain the best nutrient (N)
estimates (Castellanos et al. 2000). Soil pH was
measured with potentiometer. Nutrients (NO3,
NH4, P, Mn, Mg, K, Ca) were analyzed with the
Morgan method. Percent organic matter was
measured by the Walkey–Black method, and tex-
ture was measured following the Bouyoucos
method. Cation exchange capacity was obtained
using ammonium acetate (Castellanos et al. 2000).
At each site, observations were made on the
occurrence of different kinds of natural and
human-related disturbance, following Vázquez-
Garcı́a and Givnish (1998). These included the
numbers of fallen or damaged trees, tree stumps,
browsed or grazed plants, and intensity of grazing
(evaluated by the amount of dung, cattle foot-
prints, cattle droppings, and track livestock), as
well as the presence absence of signs of past fires,
erosion, and forest harvesting. The amount of
disturbance was summarized, with a high value
signifying high levels of disturbance and low value
signifying low levels of disturbance (Vázquez-
Garcı́a and Givnish 1998).
Data analysis
Data were summarized using four community
matrices and one environmental matrix. Two
community matrices consisted of tree importance
values and tree presence–absence data for 63 spe-
cies. Two other community matrices included
importance values and presence–absence of seed-
lings and saplings for 38 species. Matrices with
importance values were meant to relate commu-
nities in terms of their composition, structure,
distribution, and age of the nine forest stands,
while presence–absence matrices were meant to
relate communities in terms of their composition
and distribution, regardless of forest structure and
age. Thus, allowing relating environmental vari-
ables to different community aspects.
Importance values for trees were calculated
from relative tree density, frequency, and domi-
nance (basal area) (Curtis and McIntosh 1951;
Cottam and Curtis 1956). We adapted importance
values for seedlings and saplings by using height,
instead of basal area, as a measure of dominance.
The environmental matrix included quantitative
data for elevation, slope, soil nutrients, soil pH,
soil organic matter, cation exchange capacity, tree
gaps, shrub gaps, herb gaps, and disturbance and
categorical data for exposure, topography, physi-
ography, stones, and rocks.
Bray and Curtis variance-regression ordination
was used in connection with the Sørensen coeffi-
cient of similarity distance. This type of ordination
gives a complete community structure regardless
of its relationship to environmental variables and
produces clear species patterns that reflect the
environmental space the way the biotic community
interprets it (Beals 1984; McCune and Grace
2002). Thus, Bray and Curtis, like most indirect
(sociological) ordination techniques, has better
appeal than direct (environmental) ordination
techniques.
Bray and Curtis can be applied to a matrix
containing any distance measure, including non-
Euclidean semi-metrics such as Sørensen (Bray
and Curtis) distance. This is important since semi-
metrics, such as the Sørensen and Jaccard
distances, are considered robust measures of eco-
logical distance (Beals 1984; Faith et al. 1987). In
contrast, reciprocal averaging (RA) and principal
component analysis force a particular distance
(Euclidean) measure on the analyst, which pro-
duces inadequate results (Beals 1973, 1984) and
RA is effective for, but limited to, single dimen-
sional data.
Furthermore, papers that have compared Bray
and Curtis with other methods provided evidence
that Bray and Curtis may perform better than the
others (Gauch et al. 1977; Robertson 1978; Gauch
and Scrugs 1979; del Moral 1980; McCune 1994)
and produces results identical to an ordination
based on fuzzy set theory (Roberts 1986). Thus,
Bray and Curtis is considered an effective ordina-
tion technique and perhaps its only serious rival is
Multidimensional Scaling (Beals 1984; Causton
1988; Ludwig and Reynolds 1988; McCune and
Beals 1993).
Endpoints for ordination were selected by vari-
ance-regression, thereby reducing shortcomings of
the original technique. The cutoff value used for
the ordination biplot was r2=0.444, which results
in a r-value that is significantly correlated with
ordination axes. The relationship between tree,
regeneration, and the environment was evaluated
using Pearson correlations between the identified
axes of the ordination and the environmental
variables. P-values were not assigned because,

strictly speaking, the ordination scores are not
independent of each other (McCune and Grace
2002). The software used was PCORD v4.0 (mul-
tivariate analyses for ecological data) (McCune
and Mefford 1999).
Results
A total of 63 tree species representing 51 genera
and 27 families were recorded in 0.9 ha. Legumi-
nosae had the largest number of species, with 14
species. Acacia was the most speciose genus with
five species. A total of 853 individual trees were
recorded in the studied area (0.9 ha), providing an
estimate of 948 trees/ha. Importance values for
trees are shown in Appendix A. All nine sites were
considered separate in each analysis and were here
summarized in one single Appendix. A total of 38
tree species in 30 genera and 18 families were re-
corded as seedlings and saplings. The total number
of individual seedlings and saplings was 368, thus
an estimated 409 individuals/ha. Importance val-
ues for seedlings and saplings are shown in
Appendix B. All nine sites were considered sepa-
rate in each analysis and were here summarized in
one single Appendix.
Bray and Curtis variance-regression ordination
axes accounted for a substantial cumulative per-
centage of variance. For trees, the ordinations
explained more of the variation in importance
value data (70.9%) than presence–absence data
(67.1%). The opposite was true, for seedlings and
saplings, for which more variation in presence–
absence (77.4%) was explained than for impor-
tance value (62.6%).
Importance values of trees
Sites 3 and 9 were selected as endpoints for axis 1,
which extracted 32.3% of the original distance
matrix (Figure 2). This axis was explained directly
by number of shrub gaps and P concentration and
inversely by herb gaps (Table 2). Fifteen of 63
species displayed a positive correlation with this
axis and had greater importance values in areas
with greater P concentration and more shrub gaps
(Table 3). Ceiba aesculifolia (H.B.K.) Britt. and
Baker and Phenax hirtus (Sw.) Wedd. displayed a
negative correlation with this axis (Table 3).
123
Figure 2. Ordination diagram for axes 1 and 2 derived from
Bray and Curtis ordination using sites (m), importance values
of tree species, and environmental variables (vectors). Corre-
lation values for each environmental variable with the two axes
are reported in Table 2.
Sites 4 and 8 were selected as endpoints for axis
2, which extracted 20.9% of the original distance
matrix. This axis was explained inversely by ele-
vation (Figure 2, Table 2). Ten species displayed a
positive correlation with this axis and had a
greater importance value with decreasing elevation
(Table 3).
Sites 2 and 7 were selected as endpoints for axis
3 (not shown), which extracted 17.7% of the ori-
ginal distance matrix. None of the measured
environmental variables were correlated with this
axis. Heliocarpus terebinthinaceus (DC.) Hochr.
and Lysiloma microphyllum Benth. displayed a
positive correlation with this axis and decreased
with increasing importance value of Phenax hirtus
and Thouinia serrata Radlk. (Table 3).
Presence–absence of trees
Sites 8 and 3 were selected as endpoints for axis 1,
which extracted 28.6% of the original distance
matrix (Figure 3). This axis was explained directly
by slope and inversely by shrub gaps and distur-
bance (Table 2). Species that were present in sites
with a greater slope and less shrub gaps and
disturbance were Acacia riparia H.B.K. (r =
0.841), Celtis iguanea (Jacq.) Sarg. (r = 0.758),
124

Table 2. Pearson correlation (r) of environmental variables and Bray and Curtis ordination axes using importance values and pres-
ence–absence data of trees.

Variable Importance value data Presence–absence-data

Axis 1 Axis 2 Axis 3 Axis 1 Axis 2 Axis 3

Elevation 0.082 0.684 0.551 0.523 0.657 0.488

Tree gaps 0.436 0.168 0.491 0.043 0.484 0.568
Shrub gaps 0.745 0.379 0.483 0.789 0.331 0.164
Herb gaps 0.689 0.207 0.554 0.655 0.540 0.309
Disturbance 0.418 0.575 0.469 0.753 0.033 0.310
Slope 0.432 0.556 0.524 0.676 0.244 0.455
Ion exchange capacity 0.485 0.095 0.380 0.213 0.845 0.276
P 0.685 0.458 0.117 0.562 0.442 0.329
NO3 0.606 0.396 0.091 0.019 0.523 0.324
Mg 0.132 0.452 0.044 0.575 0.669 0.294
K 0.662 0.101 0.018 0.322 0.542 0.283
% Organic matter 0.232 0.288 0.025 0.475 0.188 0.229
pH 0.375 0.096 0.215 0.536 0.305 0.362

High correlations are in bold.

Conzattia multiflora (B.L. Rob.) Standl. (r = 0.735),

Table 3. Pearson correlation (r) of tree importance values and
Senna atomaria (L.) Irwin and Barneby (r = 0.726)
Bray and Curtis ordination axes.
and Spondias purpurea L. (r = 0.758). Species that
Axis 1 Axis 2 Axis 3 were absent in sites with greater slope and less shrub
Acacia cochliacantha 0.784
gaps and disturbance were Casearia corymbosa
Acacia macracantha 0.854 H.B.K. (r = 0.695), Nopalea auberi (r= 0.714),
Acacia pennatula 0.849 Senna mollisima (r= 0.695), and Stemmadenia
Adelia barbinervis 0.874 donnell-smithii (Rose) Woodson (r= 0.701).
Albizia tomentosa 0.923 Sites 9 and 1 were selected as endpoints for axis
Casearia corymbosa 0.701
Ceiba aesculifolia 0.781
2, which extracted 23.2% of the original distance
Enterolobium cyclocarpum 0.849 matrix. This axis was explained directly by ion
Exostema mexicanum 0.882 exchange and Mg concentration (Figure 3, Ta-
Ficus cotinifolia 0.882 ble 2). Acacia macilenta Rose (r = 0.697), Acaly-
Ficus insipida 0.882 pha cincta Muell. Arg. (r = 0.697), Aeschynomene
Heliocarpus terebinthinaceus 0.725
Jacaratia mexicana 0.817
amorphoides (S. Watson) Rose ex B.L. Rob.
Lasiocarpus ferrugineus 0.801 (r = 0.697), Hamelia jorullensis H.B.K.
Lysiloma microphyllum 0.768 (r = 0.697), Hintonia latiflora (Sessé and Moc. ex
Malpighia ovata 0.882 DC.) Bullock (r = 0.691), Nopalea karwinskiana
Margaritaria nobilis 0.892 (Salm-Dyck) Schumann (r = 0.758), Pachycereus
Nopalea auberi 0.791
Opuntia fuliginosa 0.913
pecten-aboriginum (Engelm.) Britt. and Rose
Phenax hirtus 0.709 0.666 (r = 0.693), Pseudobombax ellipticum (S. Watson)
Pithecellobium acatlense 0.849 Dugand (r = 0.697), and Triumfetta semitriloba
Psidium guajava 0.849 Jacq. (r = 0.697) were present in sites with higher
Senna mollisima 0.880 ion exchange and MgO; while Acacia cochliacan-
Stemmadenia donnell-smithii 0.886
Stemmadenia 0.821
tha Humb. and Bonpl. ex Willd. (r = 0.733),
tomentosa var. palmeri Heliocarpus terebinthinaceus (r= 0.697), and
Tabebuia chrysantha 0.780 Iresine cassiniformis Schauer (r= 0.669) were
Thouinia serrata 0.666 absent in sites with higher ion exchange and con-
Zanthoxylum fagara 0.849 centration Mg.
Only correlations greater than an absolute value of 0.666 are Sites 9 and 7 were selected as endpoints for axis
shown. 3, which extracted 15.3% of the original distance

Figure 3. Ordination diagram for axes 1 and 2 derived from
Bray and Curtis ordination using sites (m), presence–absence
data of tree species, and environmental variables (vectors).
Correlation values for each environmental variable with the two
axes are reported in Table 2.
matrix. None of the measured variables explained
this axis.
Importance value of seedlings and saplings
Sites 8 and 3 were selected as endpoints for axis
1, which extracted 34.3% of the original distance
matrix (Figure 4). This axis was explained in-
versely by shrub gaps (Table 4). Two species
(Ceiba aesculifolia and Lysiloma microphyllum)
displayed a positive correlation with this axis
and had greater importance values in areas with
lower number of shrub gaps. Seven species dis-
played a negative correlation with this axis
(Table 5).
Sites 7 and 9 were selected as endpoints for axis
2, which extracted 17.2% of the original distance
matrix (Figure 4). This axis was explained directly
by tree gaps and inversely by Mg and K concen-
tration (Table 4). Six species displayed a positive
correlation with this axis and had greater impor-
tance values in areas with greater number of tree
gaps and lower Mg and K concentration (Table 5).
Spondias purpurea displayed a negative correlation
125
Figure 4. Ordination diagram for axes 1 and 2 derived from
Bray and Curtis ordination using sites (m), importance values
of seedlings and saplings species, and environmental variables
(vectors). Correlation values for each environmental variable
with the two axes are reported in Table 4.
with this axis and its importance value was lower
with lower values of Mg and K (Table 5).
Sites 5 and 4 were selected as endpoints for
axis 3, which extracted 11.1% of the original
distance matrix (Figure 4). None of the measured
variables explained this axis. Acacia riparia,
Bursera simaruba (L.) Sarg., Cordia inermis
(Mill.) I.M. Johnst., and Croton fragilis H.B.K.
were positively correlated with this axis and their
importance values increased with decreasing
importance values of Bunchosia palmeri S. Wat-
son, Comocladia engleriana Loes., Senna mollis-
ima, and Tabebuia chrysantha (Jacq.) G. Nicolson
(Table 5).
Presence–absence of seedlings and saplings
Sites 8 and 4 were selected as endpoints for axis
1, which extracted 36% of the original distance
matrix. This axis was explained inversely by percent
organic matter (Figure 5). Casearia corymbosa
(r= 0.772), Nopalea auberi (r= 0.772), Senna
mollisima (r= 0.690), Spondias purpurea
(r= 0.721), and Tabebuia chrysantha (r = 0.690)
were present in sites with the most organic matter.
126

Table 4. Pearson correlation (r) of environmental variables and Bray and Curtis ordination axes using importance values and pres-
ence–absence data for seedlings and saplings.

Variable Importance value data Presence–absence-data

Axis 1 Axis 2 Axis 3 Axis 1 Axis 2 Axis 3

Elevation 0.452 0.131 0.543 0.567 0.256 0.360

Tree gaps 0.247 0.689 0.090 0.201 0.499 0.099
Shrub gaps 0.718 0.110 0.251 0.593 0.474 0.294
Herb gaps 0.587 0.248 0.275 0.393 0.642 0.427
Disturbance 0.629 0.305 0.339 0.665 0.070 0.195
Slope 0.644 0.169 0.210 0.597 0.271 0.077
Ion exchange capacity 0.307 0.527 0.141 0.070 0.807 0.341
P 0.502 0.527 0.024 0.257 0.627 0.318
NO3 0.237 0.633 0.058 0.123 0.574 0.016
Mg 0.310 0.704 0.418 0.510 0.527 0.001
K 0.288 0.689 0.238 0.064 0.712 0.515
% Organic matter 0.590 0.330 0.192 0.703 0.122 0.549
pH 0.377 0.006 0.098 0.316 0.065 0.115

High correlations are in bold.

Table 5. Pearson correlation (r) of seedlings and saplings importance values and Bray and Curtis ordination axes.

Axis 1 Axis 2 Axis 3

Acacia cochliacantha 0.798

Acacia riparia 0.750
Adelia barbinervis 0.762 0.680
Bunchosia palmeri 0.728
Bursera simaruba 0.750
Casearia corymbosa 0.699
Ceiba aesculifolia 0.679
Comocladia engleriana 0.728
Cordia inermis 0.750
Croton fragilis 0.750
Croton ciliato-glandulifera 0.798
Lasiocarpus ferrugineus 0.798
Lysiloma microphyllum 0.951
Nopalea auberi 0.821
Opuntia fuliginosa 0.732 0.726
Senna mollisima 0.737 0.680
Spondias purpurea 0.736
Stemmadenia donnell-smithii 0.897
Tabebuia chrysantha 0.673
Zanthoxylum fagara 0.798

Only correlations greater than an absolute value of 0.666 are shown.

Sites 9 and 2 were selected as endpoints for axis
2, which extracted 27.9% of the original distance
matrix. This axis was explained directly by ion
exchange and K concentration (Figure 5). Ceiba
aesculifolia (r = 0.795) and Lysiloma microphyl-
lum (r = 0.730) were present in sites with high ion
exchange and K. Acacia cochliacantha (r = 0.795),
Adelia barbinervis Schlecht. and Cham.(r = 0.730),
Croton ciliato-glandulifera Ort. (r = 0.795), Lasio-
carpus ferrugineus Gentry (r = 0.795), Opuntia
fuliginosa Griff. (r = 0.730), Senna mollisima
(r = 0.692), Stemmadenia tomentosa var. palmeri
(Rose) Woodson (r = 0.730), Tabebuia chrysantha
(r = 0.692), and Zanthoxylum fagara (L.)
C. Sargent (r = 0.795) were present in sites with
less ion exchange and K concentration.

Sites 6 and 1 were selected as endpoints for axis
3, which extracted 13.5% of the original distance
matrix. None of the measured variables explained
this axis.
Discussion
A total of 51 of the 65 species were associated
with environmental variables using Bray and
Curtis ordination and Pearson correlation. Our
data support the hypothesis that the importance
values of trees and seedlings and saplings are
related to the interaction between soil resources
and light gaps because soils in light gaps are
found to be nutrient rich, due to increased min-
eralization of organics under light conditions
(Denslow et al. 1998). In addition, presence or
absence of trees (i.e. adult stage) might be
determined by disturbance and slope, while
presence or absence of seedlings and saplings (i.e.
early life stage) might be strongly determined by
soil variables. Webb and Peart (2000), using an
ordination technique also found physiographic
and light associations in 21 of 45 species of
tropical seedlings and trees.
Importance values of trees
Availability of P, shrub gaps, and herb gaps ex-
plained most of the variation in importance values
of the overall tree community and, especially the
distribution of 17 tree species. Growth differences
in tropical trees are often found by increasing
P-supply (Chapin 1980; Burslem et al. 1994). The
greater importance value of legume, N-fixing, trees
at La Escondida-La Cabaña (40% of total number
of trees) could result from high levels of P and low
levels of N, which might enhance symbiosis, sur-
vivorship, and growth (Vitousek and Howarth
1991). Legumes trees, such as Acacia cochliacan-
tha, Acacia macracantha, Acacia pennatula
(Schlecht. and Cham.) Benth., and Enterolobium
cyclocarpum (Jacq.) Griseb., behave as light-
demanding species with a strong response in bio-
mass allocation and growth with increased P
availability, as well as a higher phosphorous-use
efficiency when P supply is low (Huante et al.
1995). In contrast, late successional (shade
tolerant) species, such as Recchia mexicana Moc.
and Sessé, have relative low growth rates and show
little or no response in growth to different P con-
centrations and less dependency on nutrient sup-
ply (Huante et al. 1995). The availability of P may
not be limiting the overall development of TDF as
shown in Chamela (Campo et al. 2001). However,
in La Escondida-La Cabaña, where slash and burn
agriculture and pasturing of livestock are common
land uses, P availability may be limiting tree spe-
cies importance value since the effects of fires and
other disturbances could increase the rate at which
P is lost from the soil (Campo et al. 2001; Louette
et al. 2001). Only Ceiba aesculifolia and Phenax
hirtus importance values seem to be favored with
decreasing P. Disturbance, together with low P,
may often limit st49mpo1-559.827598.44mbo96261t001; Louette
128
importance values of 15 species in sites with more
shrub gaps and P might be also related to high soil
resource availability that is often found in gap sites
(Denslow 1980, 1998).
The fact that elevation explained secondary axis
of importance values of the overall tree commu-
nity suggests that the effect of short elevation
gradients may not be an overriding factor on the
organization of community at small scales, where
other factors, such as nutrient supply and natural
enemies become more important (Lieberman et al.
1985; Vargas-Rodriguez 1998; Vázquez-Garcı́a
and Givnish 2000). Only rarely has elevation been
the major variable explaining community organi-
zation along short elevational gradients (Lott et al.
1987). In addition, elevation showed no relation-
ship to species richness along a larger elevational
gradient at the El Tecolote ravine, western Sierra
de Manantlán (Cuevas-G. 2002). However, the
influence of anthropogenic disturbance to this
area, may confound the effects of elevation on the
species composition of this community (Louette
et al. 2001). In our study, evergreen species such as
Ficus cotinifolia H.B.K. Ficus insipida Willd.
Albizia tomentosa (Micheli) Standl. Exostema
mexicanum A. Gray, and Margaritaria nobilis
L. increased in importance value as elevation
decreased. This may be attributed more to an
increase in soil moisture in wetter sites, close to
stream of lower areas. Changes in tree basal area,
height, and stem diameter, factors that contribute
to our estimation of importance value, were also
found to be dependent on (Segura et al. 2003).
Presence–absence of trees
Shrub gaps, slope and degree of disturbance were
the primary environmental variables that ex-
plained the variation in the presence–absence of
trees. Competition for light between subcanopy
and canopy trees in different successional stages
may be occurring, and contribute to species
diversity. For example, canopy tree Lysiloma
microphyllum formed a dense shade at one site,
preventing the establishment of pioneer species.
Consequently, gap dynamics and the interaction
of canopy and subcanopy gaps, create heteroge-
neity in light distribution throughout TDF and
can therefore influence species composition
(Montgomey and Chazdon 2001; Quigley and
Platt 2003). Anthropogenic activity also plays a
major role in the composition of La Escondida-
La Cabaña TDF forest, in which the presences of
14 tree species are related to disturbance. The
influence of disturbance on forest composition
and structure also has been found in Central and
South American TDF forest (Gonzalez and Zak
1996; Gillespie et al. 2000). Species correlations
with gaps and disturbance are often found
(Denslow 1987; Oliveira-Filho et al. 1998), but in
contrast with the present data, soil variables also
produced correlations, and explained important
variation for axis 2.
Patchy availability of nutrients in tropical dry
forest confers special patterns in microbiological
soil activity (Roy and Singh 1994). Higher
amounts of organic C, N, and P are available at
fine microsites scales and attract fine roots from
surrounding areas to support tree growth (Roy
and Singh 1994). Acacia spp. create islands of
fertility with larger soil microbial biomass, C
and N mineralization, and organic and total N
(Reyes-Reyes et al. 2002). Therefore, the patchy
availability of ion exchange and Mg may be
determining the presence of 12 tree species in La
Escondida-La Cabaña forest, which are probably
specialized to microhabitats with these soil char-
acteristics (Burslem et al. 1995; Swaine 1996).
In this study, species such as Celtis iguanea,
Conzzatia multiflora, and Senna atomaria were
associated with slopes. Studies of habitat associa-
tions in mesic to wet tropical forests also have
found slope-specialists (Clark et al. 1998; Harms
et al. 2000; Webb and Peart 2000). The spatial
distribution of soil resources may result from dis-
turbance history or differences in slope character-
istics, since gaps tend be more abundant on steep
slopes (Poorter et al. 1994). In addition, species
associated to slopes might be responding to a gra-
dient of soil characteristics such as water avail-
ability (Becker et al. 1988), nutrients (Botschek et
al. 1996; Gonzalez and Zak 1996), or soil texture
(Chauvel et al. 1987).
Importance value of seedlings and saplings
Gaps in the shrub and tree layers create hetero-
geneity of light in the understory. Not surprisingly,
importance values of seedlings and saplings were
strongly related to the presence of gaps. In high

light conditions, tree seedlings achieved higher
relative growth rates and net assimilation rates
(Rincon and Huante 1993). Pioneer species, in
particular, are light demanding and are more
negatively affected by low light environments than
shade-tolerant species (Rincon and Huante 1993).
The pioneers Lysiloma microphyllum and Ceiba
aesculifolia had lower importance values in areas
with few gaps. These results suggest that the
importance values of seedlings and saplings de-
pend on light availability. Light limitation may
reduce growth in the sapling stage, but does not
contribute to increased mortality because suscep-
tible seedlings will have already died. Seasonal
openings due to tree canopy deciduousness pro-
vide temporal sites of high light, allowing different
growth rates depending on the season (Rincon and
Huante 1993). Therefore, seedlings and saplings of
TDF species might be able to persist under a range
of light environments, with little selection for light
resource. In this sense, the classical theory of niche
specialization does not apply in seedlings and
sapling TDF populations (Welden et al. 1991;
Brokaw and Busing 2000).
Ordination axes for seedlings and saplings were
explained by K and Mg concentrations, nutrients
that play important roles in plant physiology. Mg
is a key element in chlorophyll structure and K
functions mainly as an osmoregulator and can
affect cell size. Both elements are especially critical
in the TDF community where they help prevent
mortality resulting from drought stress. For in-
stance, TDF seedlings respond to drought stress
with an increase in chlorophyll concentration
(Khurana and Singh 2001). In addition, K and Mg
concentrations appear to be correlated with tree
species richness in the tropics and are considered
as limiting resources in Amazonian forests (Gentry
1988; Burslem et al. 1995; Marschner 1995). A
response in growth with increasing Mg has been
found in tropical trees (Burslem et al. 1995;
Gunatilleke et al. 1997). Seasonal rainfall and its
effects on the reduction of microbial activity dur-
ing the dry season affect the availability of these
elements (Campo et al. 1998), and consequently,
the distribution and importance value of seedlings
and saplings. Pioneer species were positively cor-
related with this axis which is consistent with the
notion that pioneers are more dependent on
nutrient availability than shade-tolerant species
(Rincon and Huante 1994).
129
Presence–absence of seedlings and saplings
The presence and distribution of tropical decidu-
ous seedlings and saplings may be more influenced
by habitat specialization for soil resources than
light. The variance in axes from ordinations of our
presence and absence data of seedlings and sap-
lings was explained by soil variables and not by
gaps. Soil organic matter affects acidity, soil
moisture and nitrogen availability resulting in
different gradients of those factors, and thereby
determining species’ distributions. In addition, soil
organic matter contributes to cation exchange
capacity, which then controls K retention. Low
densities of seedlings between 1–30 cm tall occur in
the tropical dry forest of Western Mexico (Vargas-
Rodriguez 1998), suggesting that mortality is high
in early life stages. This is also consistent with a
higher mortality of seedlings in ‘‘suboptimal’’
habitats found in a Bornean rain forest (Webb and
Peart 2000). Poor nutrient conditions (low levels of
cation exchange capacity), or the inability of
seedlings to form symbioses with mycorrhizas
could explain limits to establishment rather than
differences in light regime resulting from canopy
gaps (Givnish 1999; Hubbell et al. 1999; Swaine
1996). Diversity in Mexican TDF may be main-
tained more by symmetric competition for soil
resources and specialization along soil microsite
gradients than by predators and pathogens related
mortality (Harms et al. 2000).
Concluding remarks
Different light conditions created by light gaps and
seasonal canopy openings influence species differ-
entiation among canopy and subcanopy trees, and
differentiation in growth and importance value of
tree populations. In contrast, seedlings and sap-
lings appear mostly to be light generalists and are
able to tolerate a wide range of light conditions
(Brokaw and Busing 2000; Wright 2002).
However, we hypothesize that habitat specializa-
tion for soil resources is likely more important in
determining species success at early stages in life
than in later stages in TDF at Jalisco, Mexico.
Also, the pattern of niche differentiation of adult
tropical trees observed along soil gradients (Clark
et al. 1999; Svenning 1999, 2001; Webb and Peart
2000) appears to occur at the seedling stage.
130

Acknowledgements Guzmán and herbaria IBUG staff helped with

This project was financed by CONACYT (96-06-002
Project), IDEAWILD and CUCBA-University of
Guadalajara. First author thanks E. Fabián
Vera Torres, Pablo Carrillo, Margarita Ayón,
Eduardo Hernández, Celso Cortés, Etelberto
Ortiz, and Ana Paula Reyes for their help with
field work. Francisco Santana Michel, Luis
species identifications. Saara Dewalt, Jennifer
Cramer, Heather Passmore, Blanca Figueroa
Rangel, and Lázaro Sánchez Velázquez made
important suggestions to this manuscript. Special
thanks Saara DeWalt for reviewing drafts of this
paper. BIOL 7093 course at Louisiana State
University provided important ideas to this
research.

Appendix A. Density, frequency, basal area, and importance value for tree species within the 0.9 ha study area in La Escondida-La
Cabaña, Jalisco, Mexico.

BA (dm2) % Freq. Density Tree/ha BA dm2/ha Sites found in I.V.

Leguminosae Acacia cochliacantha 6.14 2.22 5.56 6.82 4,9 0.45

Leguminosae Acacia macilenta 3.07 1.11 2.22 3.42 1 0.20
Leguminosae Acacia macracantha 3.97 4.44 7.78 4.41 8,9 0.64
Leguminosae Acacia pennatula 0.41 1.11 1.11 0.46 9 0.12
Leguminosae Acacia riparia 5.31 6.67 7.78 5.90 3,4 0.81
Euphorbiaceae Acalypha cincta 0.13 1.11 1.11 0.15 1 0.11
Euphorbiaceae Adelia barbinervis 2.64 2.22 2.22 2.93 8,9 0.27
Leguminosae Aeschynomene amorphoides 0.06 1.11 1.11 0.07 1 0.11
Leguminosae Albizia tomentosa 30.06 15.56 23.33 33.40 1,5,6,8 2.37
Malpighiaceae Bunchosia palmeri 13.44 5.56 7.78 14.93 2,5 0.87
Burseraceae Bursera fagaroides 6.48 5.56 5.56 7.20 3,4,5,6,9 0.67
Burseraceae Bursera grandifolia 32.94 14.44 17.78 36.61 1,2,5,7 2.15
Burseraceae Bursera kerberi 2.32 1.11 1.11 2.58 5 0.15
Burseraceae Bursera simaruba 19.29 4.44 5.56 21.44 4 0.82
Flacourtiaceae Casearia corymbosa 3.92 7.78 17.78 4.35 5,7,8,9 1.21
Bombacaceae Ceiba aesculifolia 197.69 40.00 51.11 219.66 1,2,3,4,5,6,7,8,9 7.88
Ulmaceae Celtis iguanaza 1.79 6.67 6.67 1.99 2,3,4 0.71
Cactaceae Cephalocereus alensis 4.42 2.22 2.22 4.91 3,5 0.30
Cochlospermaceae Cochlospermum vitifolium 0.14 2.22 2.22 0.16 1,2,9 0.23
Rhamnaceae Colubrina triflora 1.27 3.33 3.33 1.41 4,5,8 0.36
Anacardiaceae Comocladia engleriana 4.95 3.33 3.33 5.50 5 0.42
Leguminosae Conzattia multiflora 1.54 3.33 3.33 1.71 1,3,4 0.36
Leguminosae Coursetia glandulosa 6.95 3.33 3.33 7.72 1,3 0.46
Capparaceae Crateva palmeri 0.08 1.11 1.11 0.09 5 0.11
Euphorbiaceae Croton fragilis 0.06 1.11 1.11 0.07 4 0.11
Leguminosae Enterolobium cyclocarpum 10.43 1.11 1.11 11.59 9 0.29
Rubiaceae Exostema mexicanum 11.89 2.22 2.22 13.22 8 0.43
Moraceae Ficus cotinifolia 8.66 2.22 3.33 9.63 8 0.41
Moraceae Ficus insipida 22.80 2.22 2.22 25.33 8 0.62
Sterculiaceae Guazuma ulmifolia 11.38 4.44 5.56 12.64 4,5,7 0.69
Rubiaceae Hamelia xorullensis 0.54 1.11 1.11 0.60 1 0.12
Tiliaceae Heliocarpus terebinthinaceus 40.80 23.33 31.11 45.33 2,3,4,5,6,7,8,9 3.34
Rubiaceae Hintonia latiflora 3.04 2.22 2.22 3.38 1,2,6 0.28
Amaranthaceae Iresine cassiniiformis 0.52 4.44 4.44 0.57 3,4,9 0.46
Caricaceae Jacaratia mexicana 46.08 16.67 17.78 51.20 1,2,4,5,7,8,9 2.53
Malpighiaceae Lasiocarpus ferrugineus 43.87 13.33 27.78 48.75 2,4,9 2.62
Leguminosae Lysiloma microphyllum 538.22 74.44 341.11 598.02 1,2,3,4,5,6,7,8,9 26.27
Malpighiaceae Malpighia ovata 0.09 1.11 2.22 0.10 8 0.15
Euphorbiaceae Margaritaria nobilis 130.38 3.33 6.67 144.87 5,8 2.72
Cactaceae Nopalea Aubert 3.61 10.00 10.00 4.02 5,6,7,8,9 1.07
Cactaceae Nopalea karwinskiana 0.05 1.11 1.11 0.06 1,2 0.11
Cactaceae Opuntia fuliginosa 0.38 1.11 1.11 0.43 8,9 0.12
Cactaceae Pachycereus pecten-aboriginum 124.47 28.89 43.33 138.30 1,2,3,5,6,8 5.60
Urticaceae Phenax hirtus 144.94 55.56 101.11 161.04 1,2,3,4,5,6,7,8 9.74
 131

Appendix A. Continued.

BA (dm2) % Freq. Density Tree/ha BA dm2/ha Sites found in I.V.

Nyctaginaceae Pisonia aculeata var. aculeata 2.33 5.56 5.56 2.59 4,5 0.60
Leguminosae Pithecellobium acatlense 1.06 1.11 1.11 1.18 9 0.13
Bombacaceae Pseudobombax ellipticum 12.83 1.11 1.11 14.26 1 0.34
Myrtaceae Psidium guajava 0.06 1.11 1.11 0.06 9 0.11
Simaroubaceae Recchia mexicana 0.56 1.11 1.11 0.62 5 0.12
Leguminosae Senna atomaria 1.54 1.11 1.11 1.71 3 0.14
Leguminosae Senna mollisima 0.04 1.11 1.11 0.04 5,6,8,9 0.11
Sapotaceae Sideroxylon capiri subsp. tempisque 60.30 3.33 3.33 67.00 2,5 1.39
Anacardiaceae Spondias purpurea 165.23 43.33 66.67 183.59 1,2,3,4,5,6,7,8,9 8.08
Apocynaceae Stemmadenia donnell-smithii 4.89 13.33 20.00 5.44 1,5,6,8,9 1.67
Apocynaceae Stemmadenia tomentosa var. Palmeri 8.53 7.78 13.33 9.47 8,9 1.13
Cactaceae Stenocereus queretaroensis 4.65 3.33 3.33 5.16 2,4,8 0.42
Bignoniaceae Tabebuia chrysantha 10.34 11.11 11.11 11.49 1,2,4,5,6,8,9 1.30
Burseraceae Terebinthus acuminata 126.25 3.33 4.44 140.27 5 2.57
Sapindaceae Thouinia serrata 16.59 4.44 5.56 18.43 1,2,5,8 0.78
Sapindaceae Thouinidium decandrum 5.62 4.44 5.56 6.24 2,5,6,8 0.59
Tiliaceae Triumfetta semitriloba 0.38 1.11 1.11 0.43 1 0.12
Rutaceae Zanthoxylum caribaeum 2.41 6.67 8.89 2.68 4,5,6,7 0.79
Rutaceae Zanthoxylum fagara 0.12 1.11 1.11 0.14 9 0.11

Totals 1914.98 947.78 2127.76

All nine sites were considered separate in each analysis and were here summarized in one single Appendix.

Appendix B. Density, frequency, mean height and importance value for seedlings and saplings species within the 0.9 ha in La Es-
condida-La Cabaña, Jalisco, Mexico.

% Freq. Density Trees/ha Mean height Sites found in I.V.

Leguminosae Acacia cochliacantha 3.33 13.33 0.53 9 2.24

Leguminosae Acacia macracantha 1.11 1.11 3.00 8 4.28
Leguminosae Acacia riparia 1.11 1.11 1.66 4 2.47
Euphorbiaceae Adelia barbinervis 8.89 12.22 0.44 8,9 2.76
Leguminosae Aeschynomene amorphoides 1.11 5.56 0.10 1 0.73
Leguminosae Albizia tomentosa 2.22 2.22 0.85 1,5 1.62
Malpighiaceae Bunchosia palmeri 2.22 2.22 1.29 5 2.21
Burseraceae Bursera fagaroides 4.44 4.44 0.45 4,6,9 1.55
Burseraceae Bursera grandifolia 17.78 33.33 0.09 1,2,7,8 5.18
Burseraceae Bursera simaruba 1.11 1.11 0.34 4 0.70
Flacourtiaceae Casearia corymbosa 21.11 41.11 0.22 5,7,8,9 6.42
Bombacaceae Ceiba aesculifolia 27.78 37.78 0.14 1,2,3,4,5,6,7,8 6.92
Cactaceae Cephalocereus alensis 4.44 5.56 0.15 3,5 1.24
Cochlospermaceae Cochlospermum vitifolium 2.22 2.22 0.42 1,2 1.04
Anacardiaceae Comocladia engleriana 1.11 1.11 0.62 5 1.07
Boraginaceae Cordia inermes 2.22 2.22 1.24 4 2.15
Leguminosae Coursetia glandulosa 2.22 5.56 0.30 3 1.15
Euphorbiaceae Croton fragilis 1.11 4.44 0.96 4 1.80
Euphorbiaceae Croton ciliato-glandulifera 6.67 11.11 0.38 9 2.29
Tiliaceae Heliocarpus terebinthinaceus 6.67 6.67 0.54 2,6,9 2.15
Rubiaceae Hintonia latiflora 1.11 1.11 1.54 1 2.31
Caricaceae Jacaratia mexicana 4.44 4.44 0.66 1,4,9 1.83
Malpighiaceae Lasiocarpus ferrugineus 6.67 8.89 0.51 9 2.28
Leguminosae Lysiloma microphyllum 28.89 53.33 0.09 1,2,3,4,5,6,7 8.27
Euphorbiaceae Margaritaria nobilis 1.11 1.11 2.50 8 3.61
Cactaceae Nopalea Aubert 11.11 28.89 0.19 5,7,8,9 4.08
Cactaceae Nopalea karwinskiana 2.22 2.22 0.59 1,2 1.27
132

Appendix B. Continued.

% Freq. Density Trees/ha Mean height Sites found in I.V.

Cactaceae Opuntia fuliginosa 3.33 3.33 0.43 8,9 1.29
Cactaceae Pachycereus pecten-aboriginum 3.33 3.33 0.33 2,5,8 1.15
Urticaceae Phenax hirtus 5.56 6.67 0.51 1,7,8 1.96
Leguminosae Senna mollisima 7.78 10 0.36 5,8,9 2.32
Anacardiaceae Spondias purpurea 15.56 23.33 0.24 1,2,3,4,5,7,8 4.47
Apocynaceae Stemmadenia donnell-smithii 25.56 45.56 0.18 1,5,8 7.32
Apocynaceae Stemmadenia tomentosa var. Palmeri 2.22 2.22 0.90 8,9 1.68
Bignoniaceae Tabebuia chrysantha 3.33 3.33 0.55 5,8,9 1.45
Sapindaceae Thouinidium decandrum 2.22 3.33 0.49 2,8 1.22
Rutaceae Zanthoxylum caribaeum 7.78 10 0.20 5,6,7 2.11
Rutaceae Zanthoxylum fagara 2.22 3.33 0.63 9 1.41
Totals 408.89

All nine sites were considered separate in each analysis and were here summarized in one single Appendix.

References Castellanos J.Z., Uvalle-Bueno J.X. and Aguilar-Santelises A.

Balvanera P., Lott E., Segura G., Siebe C. and Islas A. 2002.
Patterns of b-diversity in a Mexican tropical dry forest. J.
Veg. Sci. 13: 145–158.
Basnet K. 1992. Effect of topography on the pattern of trees in
Tabonuco (Dacryodes-Excelsa) dominated rain-forest of
Puerto-Rico. Biotropica 24: 31–42.
Battles J.J., Dushoff J.G. and Fahey T.J. 1996. Line intersect
sampling of forest canopy gaps. Forest Sci. 42: 131–138.
Beals E.W. 1973. Ordination: mathematical elegance and eco-
logical naivete. J. Ecol. 61: 23–35.
Beals E.W. 1984. Bray-Curtis Ordination: an effective strategy
for analysis of multivariate ecological data. Adv. Ecol. Res.
14: 1–55.
Becker P., Rabenold P.E., Idol J.R. and Smith A.P. 1988.
Water potential gradients for gaps and slopes in a Panama-
nian tropical moist forest’s dry season. J. Trop. Ecol. 4: 173–
184.
Botschek J., Ferraz J., Jahnel M. and Skowronek A. 1996. Soil
chemical properties of a toposequence under primary forest
in the Itacoatiara vicinity (Amazonas, Brazil). Geoderma 72:
119–132.
Brokaw N. and Busing R.T. 2000. Niche versus chance and tree
diversity in forest gaps. Tree 15: 183–188.
Burslem D.F.R.P., Turner I.M. and Grubb P.J. 1994. Mineral
nutrient status of coastal hill dipterocarp forest and adinan-
dra belukar in Singapore: bioassays of nutrient limitation. J.
Trop. Ecol. 10: 579–600.
Burslem D.F.R.P., Turner I.M. and Grubb P.J. 1995. Re-
sponses to nutrient addition among shade-tolerant tree
seedlings of lowland rain forest in Singapore. J. Ecol. 83:
113–122.
Campo J., Jaramillo V.J. and Mass J.M. 1998. Pulses of soil
phosphorus availability in a Mexican tropical dry forest: ef-
fects of seasonality and level of wetting. Oecologia 115: 167–
172.
Campo J., Mass M., Jaramillo V.J., Martinez-Yrizar A. and
Sarukhan J. 2001. Phosphorus cycling in a Mexican tropical
dry forest ecosystem. Biogeochemistry 53: 161–179.
2000. Memoria del curso sobre interpretación de análisis de
suelos, aguas agrı́colas y plantas. Instituto Nacional de
Investigaciones Forestales, Agrı́colas y Pecuarias, México.
Causton D.R. 1988. An Introduction to Vegetation Analysis:
Principles, Practice and Interpretation. London, UK.
CETENAL 1975. Carta geológica, escala 1:50 000. Insti-
tuto Nacional de Estadı́stica, Geografı́a e Informática,
México.
CETENAL 1976. Carta edafológica, escala 1:50 000. Instituto
Nacional de Estadı́stica, Geografı́a e Informática, México.
Chapin F.S.III 1980. The mineral nutrition of wild plants.
Annl. Rev. Ecol. Sys. 11: 233–260.
Chauvel A., Lucas Y. and Boulet R. 1987. On the genesis of the
soil mantle of the region of Manaus, Central Amazonia,
Brazil. Experientia 43: 234–241.
Clark D.B., Clark D.A. and Rich P.M. 1993. Comparative
analysis of mirohabitat utilization by saplings of nine tree
species in neotropical rainforest. Biotropica 25: 397–407.
Clark D.B., Clark D.A. and Read J.M. 1998. Edaphic variation
and the mesoscale distribution of tree species in a neotropical
rain forest. J. Ecol. 86: 101–112.
Clark D.B., Palmer M.W. and Clark D.A. 1999. Edaphic fac-
tors and the landscape-scale distributions of tropical rain
forest trees. Ecology 80: 101–112.
Connell J. 1978. Diversity in tropical rain forest and coral reefs.
Science 299: 1302–1310.
Cottam G. and Curtis J.T. 1956. The use of distance measures
in phytosociological sampling. Ecology 37: 451–460.
Cruz P. 1989. Atlas cartográfico de la reserva de la biosfera
Sierra de Manantlán, Jalisco, México. Laboratorio Natural
Las Joyas, Universidad de Guadalajara, México.
Cuevas G.R. 2002. Análisis de gradientes de la vegetación de la
cañada El Tecolote, en la Sierra de Manantlán, Jalisco,
México. PhD. Dissertation, Colegio de Postgraduados,
Montecillo, México.
Cuevas G.R., Nuñez L.N., Guzman H.L. and Santana M.F. 1998.
El bosque tropical caducifolio en la reserva de la biosfera Sierra
de Manantlán, Jalisco-Colima, México. Boletı́n del Instituto de
Botánica de la Universidad de Guadalajara 5: 445–491.
Curtis J.T. and McIntosh R.P. 1951. An upland forest contin-
uum in the prairie forest border region of Wisconsin. Ecology
32: 476–496.
del Moral R. 1980. On selecting indirect ordination methods.
Vegetatio 42: 75–84.
Denslow J.S. 1980. Gap partitioning among tropical rainforest
trees. Biotropica 12(suppl.): 47–55.
Denslow J.S. 1987. Tropical rain forest gaps and tree species
diversity. Annl Rev. Ecol. Sys. 18: 431–451.
Denslow J.S., Ellison A.M. and Sanford R.E. 1998. Tree fall
gap size effects on above- and below-ground processes in a
tropical wet forest. J. Ecol. 86: 597–609.
Faith D.P., Minchin P.R. and Belbin L. 1987. Compositional
dissimilarity as a robust measure of ecological distance.
Vegetatio 69: 57–68.
Gauch H.G.Jr. and Scruggs W.M. 1979. Variants of polar
ordination. Vegetatio 40: 147–153.
Gauch H.G.Jr., Whittaker R.H. and Wentworth T.R. 1977. A
comparative study of reciprocal averaging and other ordi-
nation techniques. J. Ecol. 65: 157–174.
Gentry A.H. 1982. Patterns of neotropical plant species diver-
sity. Evol. Biol. 15: 1–84.
Gentry A.H. 1988. Changes in plant community diversity and
floristic composition on environmental and geographical
gradients. Annl. Missouri Bot. Garden 75: 1–34.
Gillespie T.W., Grijalva A. and Farris C.N. 2000. Diversity,
composition and structure of tropical dry forest in Central
America. Plant Ecol. 147: 37–47.
Givnish T.J. 1999. On the causes of gradients in tropical tree
diversity. J. Ecol. 87: 193–210.
Gonzalez O.J. and Zak D.R. 1996. Tropical dry forest of St.
Lucia, West Indies: Vegetation and soil properties. Biotro-
pica 28: 618–626.
Gunatilleke C.V.S., Gunatilleke I.A.U.N., Perera G.A.D.,
Burslem D.F.R.P., Ashton P.M.S. and Ashton P.S. 1997.
Responses to nutrient addition among seedlings of eight
closely related species of Shorea in Sri Lanka. J. Ecol. 85:
301–311.
134
Murphy P. and Lugo A. 1986. Structure and biomass of a
subtropical dry forest in Puerto Rico. Biotropica 2: 89–96.
Oliveira-Filho A.T., Curi N., Vilela E.A. and Carvalho D.A.
1998. Effects of canopy gaps, topography, and soils on the
distribution of woody species in a central Brazilian deciduous
dry forest. Biotropica 30: 362–375.
Olvera M., Moreno S. and Figueroa B. 1996. Sitios Perma-
nentes para la investigación silvı́cola, manual para su estab-
lecimiento. Universidad de Guadalajara, México.
Pearson T.R., Burslem D.F.R.P., Goeriz R.E. and Dalling J.W.
2003. Interactions of gap size and herbivory on establish-
ment, growth and survival of three species of neotropical
pioneer trees. J. Ecol. 91: 785–796.
Phillips O. and Miller J.S. 2002. Global patterns of plant
diversity: Alwyn H. Gentry’s forest transect data set. Mis-
souri Botanical Garden Press, Missouri, US, pp. 319.
Poorter L., Jans L., Bongers F. and van Rompaey R.S.A.R.
1994. Spatial distribution of gaps along three catenas in the
moist forest of Tai National Park, Ivory Coast. J. Trop. Ecol.
10: 385–398.
Poulsen A.D. and Baslev H. 1991. Abundance and cover of
ground herbs in an Amazonian rainforest. J. Veg. Sci. 2: 315–
322.
Quigley M.F. and Platt W.J. 2003. Composition and structure
of seasonally deciduous forest in the Americas. Ecol. Mo-
nogr. 73: 87–106.
Reyes-Reyes G., Baron-Ocampo L., Cuali-Alvarez I., Frias-
Hernandez J.T., Olalde-Portugal V., Fregoso L.V. and
Dendooven L. 2002. C and N dynamics in soil from the
central highlands of México as affected by mesquite (Prosopis
spp.) and huizache (Acacia tortuoso): a laboratory investi-
gation. Appl. Soil Ecol. 19: 27–34.
Rincon E. and Huante P. 1993. Growth responses of tropical
deciduous tree seedlings to contrasting light conditions. Trees
7: 202–207.
Rincon E. and Huante P. 1994. Influence of mineral nutrient
availability on growth of tree seedlings from the tropical
deciduous forest. Trees 9: 93–97.
Roberts D.W. 1986. Ordination on the basis of fuzzy set theory.
Vegetatio 66: 123–131.
Robertson P.A. 1978. Comparison of techniques for ordinating
and classifying old-growth floodplain forest in southern Illi-
nois. Vegetatio 37: 43–51.
Roy S. and Singh J.S. 1994. Consequences of habitat hetero-
geneity for availability of nutrients in a dry tropical forest. J.
Ecol. 82: 503–509.
Rzedowski J. 1978. Vegetación de México. Editorial Limusa,
México.
Segura G., Balvanera P., Durán E. and Pérez A. 2003. Tree
community structure and stem mortality along a water
availability gradient in a Mexican tropical dry forest. Plant
Ecol. 169: 259–271.
Svenning J.C. 1999. Microhabitat specialization in a species-rich
palm community in Amazonian Ecuador. J. Ecol. 87: 55–65.
Svenning J.C. 2001. Environmental heterogeneity, recruitment
limitation and the mesoscale distribution of palms in a
tropical montane rain forest (Maquipucuna, Ecuador). J.
Trop. Ecol. 17: 97–113.
Swaine M.D. 1996. Rainfall and soil fertility as factors limiting
forest species distributions in Ghana. J. Ecol. 84: 419–428.
Terborgh J. 1973. On the notion of favorableness in plant
ecology. Am. Nat. 107: 481–501.
Trejo I. 1998. Distribución y diversidad de las selvas bajas de
México: relaciones con el clima y suelo. PhD. Dissertation,
Universidad Nacional Autónoma de México, México, DF.
Trejo I. and Dirzo R. 2000. Deforestation of seasonally dry
tropical forest: a national and local analysis in Mexico. Biol.
Conserv. 94: 133–142.
Trejo I. and Dirzo R. 2002. Floristic diversity of Mexican
seasonally dry tropical forests. Biodiv. Conserv. 11: 2063–
2048.
Tuomisto H. and Ruokolainen K. 1993. Distribution of Pter-
idophyta and Melastomataceae along an edaphic gradient in
an Amazonian rain-forest. J. Veg. Sci. 5: 25–34.
Vargas-Rodriguez Y.L. 1998.Odenación sociológica de la co-
munidad arbórea del bosque tropical caducifolio en El Ag-
uacate-Zenzontla, Sierra de Manantlán, Jalisco. Bc.S. thesis,
Centro Universitario de Ciencias Biológico Agropecuarias,
Universidad de Guadalajara, Guadalajara.
Vázquez-Garcı́a J.A., Cuevas G. R., Cochrane T.S., Iltis H.H.,
Santana M.F.J. and Guzman H.L. 1995. Flora de Mana-
ntlán. Sida, Botanical Miscellany 13. Botanical Research
Institute of Texas, USA.
Vázquez-Garcı́a J.A. and Givnish T.J. 1998. Altitudinal gradi-
ents in tropical forest composition, structure, and diversity in
the Sierra de Manantlán. J. Ecol. 86: 999–1020.
Vázquez-Garcı́a J.A. and Givnish T.J. 2000. Vegetation of the
cerro Grande massif, Sierra de Manantlan, Mexico: ordina-
tion of a long altitudinal gradient with high species turnover.
Boletı́n del Instituto de Botánica 6: 227–250.
Vitousek P.M and Howarth R.W. 1991. Nitrogen limitation on
land and in the sea: how can it occur? Biogeochemistry 13:
87–115.
Webb C.O. and Peart D.R. 2000. Habitat associations of trees
and seedlings in a Bornean rain forest. J. Ecol. 88: 464–478.
Welden C.W., Hewett S.W., Hubbell S.P. and Foster R.B. 1991.
Sapling survival, growth, and recruitment: relationship to
canopy eight in a neotropical forest. Ecology 72: 35–50.
Whittaker R.H. 1956. Vegetation of the Great Smoky Moun-
tains. Ecol. Monogr. 26: 1–80.
Whittaker R.H. 1960. Vegetation of the Siskiyou Mountains,
Oregon and California. Ecol. Monogr. 30: 279–338.
Wright S.J. 2002. Plant diversity in tropical forest: a review of
mechanisms of species coexistence. Oecologia 130: 1–14.