POLISH JOURNAL OF ECOLOGY

(Pol. J. Ecol.)

61

283295

2013

Regular research paper

Pablo MARTNEZ-ANTNEZ 1, Christian WEHENKEL 2, J. Ciro HERNNDEZ-DAZ 2 *,

Martha GONZLEZ-ELIZONDO 3, J. Javier CORRAL-RIVAS 4, Alfredo PINEDO-LVAREZ 2
Doctorado Institucional en Ciencias Agropecuarias y Forestales, Universidad Jurez del Estado de
Durango, Km 5.5 Carretera Mazatln, 34120 Durango, Mxico
2
Instituto de Silvicultura e Industria de la Madera, Universidad Jurez del Estado de Durango, Km 5.5
Carretera Mazatln, 34120 Durango, Mxico; *e-mail: jciroh@ujed.mx (corresponding author)
3
Instituto Politecnico National, CIIDIR, Unidad Durango. Sigma 119 Col. 20 de Noviembre II.
Durango, 34220 Mxico
4
Facultad de Ciencias Forestales, Universidad Jurez del Estado de Durango, Ro Papaloapan y Blvd.
Durango s/n, Col. Valle del Sur, 34120 Durango, Mxico
1

EFFECT OF CLIMATE AND PHYSIOGRAPHY ON THE DENSITY

OF TREE AND SHRUB SPECIES IN NORTHWEST MEXICO
ABSTRACT: In order to understand the environmental variables that may impact more on
the distribution of species of trees and shrubs, a
correlation analysis applying the Covariation (C)
of Gregorius was conducted among 14 variables of
climate and physiography, and the number of individuals of 72 species, which were found in 1804
sampling plots (covering about 123,317 km2) of
the National Forests and Soils Inventory (INFyS)
developed by the National Forest Commission in
Mexico (CONAFOR). Among the studied species
there are several of the genera Quercus, Pinus and
Junniperus, which are mainly distributed in the Sierra Madre Occidental, where they stand out for
their abundance.
The results show that the density of 88% of the
studied species have a significant correlation (P
<0.025) with at least five of the 14 variables analyzed. Seven of the variables showed significant correlation (P <0.025) with at least 74% of the studied
species: Julian date of last spring frost with an average value of covariation (C) equal to 0.71, average
duration of the frost-free period with average value
of C = 0.71, degree days above 5oC with covariation
of 0.69, altitude above sea level with C = 0.66; mean
temperature in the coldest month, mean temperature in the warmest month and mean annual temperature, with average values of C = 0.65 for each
of these last three variables. The geographic orientation of the ground was the least correlated with
the density of the species, since only 10% of them
showed significant correlation with this variable.

journal 34.indb 283

KEY WORDS: Mexican dendroflora, plant

species adaptation, bioclimatic niche
1. INTRODUCTION
A variety of physiographic and site specific factors influence the presence of plant
species (Wo o dw ard et al. 2004), including:
land relief, soil depth, altitude above sea level
and geographic location (Wo o dw ard 1987,
Ste f fe n 2008).
The characteristics of climate often become decisive factors for the presence of
plants (B e g et al. 2002, Steffen 2008, S e n z R omero et al. 2010). Some of the climate
variables that affect the growth of plants are:
temperature, sunshine duration, season of the
year, length of the growth period, and others
(Ko uchowsk i and Teg i r mend i 2005).
The study of climate of the last two million years has shown that the abiotic environment remains in a state of non-equilibrium,
and environmental conditions follow changing trends with periods of different lengths
(R eh feldt 2006, R eh feldt et al. 2006).
However, plant species have not shown a significant morphological response to climate
change, but instead there is evidence of dispersion, population fragmentation and even

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Pablo Martinez-Antnez et al.

extinction (Hug hes 2000, Peters on and

Vieg l ais 2001, G op araju and Jha 2010).
Recent research suggests that there has
been a change in the global climate system.
Some of these studies document changes in
environmental conditions over short periods
of time, threatening the survival of ecosystems
and species (IPCC-WGI 2007, Pachau r i
and Ja l low 2007, Senz-R omero et al.
2010, C onde et al. 2011).
Several authors agree that the increase in
global temperature affect biodiversity at different scales and ways, geographically shifting plant and animal communities and altering the ecological niche of organisms and the
ecosystem function (Desai et al. 2008, Zhu
et al. 2011).
It has been stated that trees and shrubs,
besides fulfilling ecological functions such
as habitat for wildlife reproduction and feeding, also have economic and social importance. For example, in forest regions forestry
is a major economic activity for many people (Nvar 2004), while in non-forest areas
many shrub and tree species are important
for medicinal purposes, protection against
soil erosion, retention of water resources,
commercial or other scientific purposes
(G on z le z-E l izondo et al. 2004, G op araju and Jha 2010).
Vegetation studies have been conducted
from several approaches, including: diversity,
frequency, distribution and density of species.
The frequency of occurrence of a species has
been measured as the percentage of incidence
of the species in a given number of sites, Maurer et al. (2003) found that more frequent species are also more abundant. Density of a species is defined as the number of individuals per
unit area (Kronenfeld and Wang 2007).
The aim of this work was to study the
degree of correlation between the density of
72 species of trees and tall shrubs with 14 climatic and physiographic variables considered
important for plants (R ehfeldt et al. 2006).
The study explores relationships between tree
species density to climatic variables such as
accumulated precipitation in the growing season, mean temperature in the warmest or the
coldest month, and date of last spring frost,
which are seldom used for inferring presence
of plant species. This knowledge may be helpful to define eco-physiographic areas useful to

journal 34.indb 284

face a possible drastic environmental change

and to strengthen other technical or scientific
purposes (Tcheb a kov a et al. 2005, Ait ke n
et al. 2008, Z hu et al. 2011).
2. MATERIALS AND METHODS
The study was conducted in the state of
Durango, which has an approximate area of
123,317 km2, 40% of the State has a dry and
semi-dry climate, 34% sub humid-temperate,
14% very dry, 11% sub humid-warm and only
1% humid-temperate (INEGI 2012). Durango
is located to the northwest of Mexico between
2653 and 2216 North and between 10229
and 10716 West (Fig. 1). In the Sierra Madre
Occidental mountain range that crosses the
State there are woodlands of Pinus, PinusQuercus, Quercus-Pinus, small portions of
temperate mesophytic forest; and on the western flanks some tropical deciduous and semi
deciduous forests (G onz le z-E lizondo
et al. 2007). In the region of the valleys there
are mainly desert and semi-desert zones. Many
of the forests in the study area, especially where
Pinus is the dominant genus, are subject to timber harvesting and, in some areas forest fires
occur in periods of drought (Wehen kel et al.
2011a, G onz le z-E lizondo et al. 2012b).
The 1804 primary sampling units (conglomerates) utilized in this study were established in most of the State by the National
Forest Commission (CONAFOR 2009), to
perform the National Inventory of Forests
and Soils (INFyS 20042009). The name of
conglomerate was given because each of
them is integrated by a group of four circular
secondary sampling units of 400 m2 in size,
known as sites. The sites were distributed
into the conglomerate with an equidistance of
45.14 m; the first site was located in the centre
of the conglomerate, the second to the north
at Az 0, the third to the southeast direction
Az 120 and the fourth to the southwest Az
240. However, the data of the INFyS is mainly reported at the level of conglomerate; so,
in the remaining of this report the word plot
is utilized instead of conglomerate; therefore
the area of each plot is 1600 m2. The network
of plots is distributed every 5 km in forested
areas of the Sierra Madre Occidental, and
every 20 km in arid and semiarid regions
(CONAFOR 2009).

2013-07-09 14:47:33

Effect of climate and physiography on the density of tree and shrub species

In each plot, all the stems of trees and

shrubs, whose diameter was greater than or
equal to 7.5 cm at the height of 1.3 m above
the ground level were counted and identified
at the species level. Following Krone n feld
and Wang (2007), the number of stems recorded in each plot is treated in this paper
as density. The whole list of studied species
is presented in Appendix I, showing the frequency of occurrence (Maurer et al. 2003)
and the mean density of each species per hectare in the study area. For example, Pinus herrerae presented a frequency of 6.04% (since it
was found in 108 out of the 1804 plots) and a
mean density of 4.76 stems per hectare.
A total of 329 species of trees and shrubs
were found in the 1804 studied plots, from
which 85 were selected for further analyses,
following the criterion of finding at least 50
individuals of each species in the 1804 plots.
Nevertheless, 13 species were later discarded
because their taxonomic identification was
very doubtful. Thus, the final data set was
composed of 72 species of trees and shrubs
(Appendix I). Over 80% of these species were
found in the plots located on the Sierra Madre
Occidental mountain range.
Four physiographic and site specific variables considered in this work include: the average slope of the terrain (AST, in %), altitude

285

above sea level (ASL, m), dominant aspect or

geographic orientation of the ground (ASP:
zenith, north, south, east, west, north-east,
south-east, north-west and south-west) and
the soil depth (SD, in cm) (CONAFOR 2009).
In addition, 10 climate variables considered
important according to the algorithms of R e h fel dt (2006), were obtained from the website of the Forest Service of the Department of
Agriculture in the United States (http://forest.
moscowfsl.wsu.edu/climate/) which models
climate data from 4000 weather stations of
Mexico, South of the United States of America, Guatemala, Belize and Cuba, with records
from 1976 to 1990 (Cro ok ston et al. 2008,
S en z - R omero et al. 2010). These data are
modelled using the ANUSPLIN software, developed by Hutch i ns on (2004).
The climate variables obtained from that
source for each of the 1804 INFyS plots, were:
mean annual temperature (MAT, C), mean
annual precipitation (MAP, mm), total precipitation in the growing season (April to
September (GSP, mm), mean temperature
in the coldest month (MTCM, C), mean
minimum temperature in the coldest month
(January) (MMIN, C), mean temperature
in the warmest month (June) (MTWM, C),
mean maximum temperature in the warmest
month (June) (MMAX, C), average length

Table 1. Descriptive statistics of the independent variables*.

Variable
Units
Minimum
Maximum
Mean
S.D.
ASP
*
0
315
128.27
109.82
ASL
m
206
4502
2219
512.52
AST
%
0
98
25
16.48
SD
cm
0
110
38
23.99
MAT
C
3.9
26.1
14
3.85
MAP
mm
250
1444
878
251.84
GSP
mm
190
1068
667
167.9
MTCM
C
1.2
21.2
8.53
3.94
MMIN
C
-7.5
11.5
-0.67
4.47
MTWM
C
6.3
30.7
18.47
3.91
MMAX
C
12.8
40.1
26.9
3.61
SDAY
days
0
196
109.07
48.91
FFP
days
43
365
196.06
75.49
DD5
C
16
253
109.89
44.35
* S.D.: Standard deviation, ASP: Geographical aspect (zenith, north, south, east, west, north-east, south-east, northwest and south-west), ASL: Altitude above sea level, AST: Average slope of the terrain, SD: Depth of the soil, MAT:
Mean annual temperature, MAP: Mean annual precipitation, GSP: Total precipitation in the growing season (April
to September), MTCM: Mean temperature in the coldest month, MMIN: Mean minimum temperature in the coldest month, MTWM: Mean temperature in the warmest month, MMAX: Mean maximum temperature in the warmest month, SDAY: Day of the year in which it is probable to happen the last frost in spring, FFP: Frost free period,
DD5: Degree days above 5C.

journal 34.indb 285

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Pablo Martinez-Antnez et al.

ranges between 1 and +1, where C = 1 corresponds to an entirely positive covariation and
C = 1 to a strictly negative covariation. If its
denominator is zero, C is undefined (Gre gor ius et al. 2007). Formally:

Fig. 1. Location of the study area.

of frost-free period (FFP, days), Julian date

of the last frost spring (SDAY, days) and degree-days above 5C (DD5, C). All of these
variables are considered critical to the full
development of plants (Tcheb a kova et al.
2005, R ehfeldt et al. 2006). Table 1 provides
a summary of the conditions prevailing in the
study area, in terms of the basic values of the
variables.
To evaluate the correlation between the
density of species and ASP (which is not a
numerical variable) using the same method,
each aspect was assigned a numerical value
indicating the predominant azimuth: Zenithal 0, North 1, North East 45, East 90, South
East 135, South 180, South West 225, West
270, and North West 315.
Given that there was not found linearity or a normal distribution of the data, the
relationship between number of individuals
per species and plot (species density) and climatic and physiographic variables was measured by the covariation (C) described by
Gre gor ius et al. (2007). By definition, two
ordinal variables X and Y show entire covariation if one variable consistently increases or
consistently decreases as the other variable
increases. There thus exists a strictly monotonic relationship (but not necessarily linear)
between the two variables. The covariation C

journal 34.indb 286

(1)

where: C = Covariation between each

species X and variable Y. (Xi-Xj) = difference
in climate values of the specie X, between the
i-th and the j-th plots. (Yi-Yj) = difference in
density value of the variable Y, between the
i-th and j-th plots.
The denominator of Eq. (1) is the summation of absolute values of the indicated
products.
In order to test the probability that the observed degrees of covariation C are produced
solely by random events rather than directed
forces, a two-sided permutation test was performed based on randomly chosen reassignments (Man ly 1997). The percentages of imitated C greater than or equal to the respective
observed C (P-values) were computed for a
satisfactory number of permutations.
3. RESULTS AND DISCUSSION
The estimated covariation values of
Gregorius (C) showed that 88% of the studied species have a significant correlation
(P <0.025) with at least five variables of climate and physiography (Appendixes II, III
and IV). The species Abutilon sp., Guazuma
ulmifolia, and Tilia mexicana presented the
highest correlations simultaneously with several variables and also, the largest mean values of C (mean C).
The climatic variables that apparently
largely affect the density of most of the studied
species are: SDAY with an average C = 0.71;
FFP with C = 0.71; DD5 with C = 0.69; ASL
with C = 0.66, and MTCM, MTWM, MAT
with average values of C = 0.65 each of these
three variables.
Analyzing by species group, conifers
(Appendix II) and Quercus (Appendix III)
showed little correlation with AST, indicating
that they may equally be found in varying degrees of slope. Some species of other genera
different than Quercus or conifers (Appendix

2013-07-09 14:47:33

Effect of climate and physiography on the density of tree and shrub species

IV) were significantly (P <0.025) correlated

with AST, showing C values between 0.62 and
0.73; they include: two shrubs (Colubrina heteroneura and Caesalpinia pulcherrima), and
two trees (Tilia mexican and Bursera coyucensis ) of tropical forest, and a temperate forest tree (Prunus serotina). Two other tropical
shrubs (Abutilon glauca and Erythrina sp.)
showed C of 0.83 and 0.85 respectively with
AST, but these values were not significant.
The ASP does not seem to be a limiting
factor for the habitat of most species, since it
is the variable less correlated with the density of the 72 species analyzed. Only seven of
them showed significant correlation with this
variable. Out of them, three are of the genus
Quercus, which is distributed entirely in the
mountainous region of the State.
Also in regard to SD, it was found no evidence of a large effect on the density of the
studied species, Juniperus monticola being the
only one showing a significant high C value
(0.90).
On the other hand it is observed that
MAT shows significant correlations with
71.4% of the group of conifers (Appendix II),
with 51.6% of the group of Quercus (Appendix III) and with 85.0% of the group of other
genera (Appendix IV). In the same order, the
percentages of absolute values of C equal to or
greater than 0.90 were respectively 9.5% for
conifers (Cupressus lusitanica and Pinus herrerae), 6.4% for Quercus (Q. tarahumara and
Q. magnoliifolia) and 55.0% for other genres
(Guazuma ulmifolia, Tilia mexicana and Colubrina heteroneura, among others), which
means that the latter group is more demanding in terms of the specific temperature requirements for their existence, implying that
their respective habitats are smaller.
The above is corroborated by noting that
similar correlations to those presented by the
three groups of species for MAT, were also
observed with respect to MTCM, MMIN,
MTWM and MMAX. It was found that some
of these variables are inter-correlated, which
was also observed by Silva-Flores and Wehenkel (unpublished); however, it was decided to
report about all of them, since one of the contributions of this work is that, in Appendixes
II, III and IV it can be seen which of these are
the variables that individually have more (or
less) influence on the density of each species.

journal 34.indb 287

287

Pines and oaks showed less requirements

with respect to precipitation that with regard to
temperature, as the observed values of C for the
variables GSP and MAP were lower than those
observed for the variables listed in the previous
paragraph. Out of the 21 species of conifers only
one (Abies durangensis), and only three out of
the 31 oaks (Quercus resinosa, Q. acutifolia and
Q. urbanii) reached C values higher than 0.90
with respect to these two variables. But, among
the other genres (Appendix IV) the effect of
precipitation was more remarkable, since three
of the six shrubs (Abutilon sp., Erythrina glauca
and Acacia berlandieri) and four of the 14 trees
(Guazuma ulmifolia, Alnus jorullensis, A. acuminata and A. firmifolia) showed values of C
greater than 0.90, which indicates that they have
specific requirements of precipitation.
Variables ASL, SDAY, FFP and DD5 seem
to have the same pattern of behaviour, considering the absolute value of C in most of
the analyzed species. Some correlations were
negative and others positive.
As happened with the variables that directly indicate temperature (MAT, MTCM,
MMIN, MTWM and MMAX) and those related to precipitation (MAP and GSP) again,
the stronger effect of ASL, SDAY, FFP and
DD5 was found in the species listed in Appendix IV, where five tropical forest shrubs (Abutilon sp., Colubrina heteroneura, Caesalpinia
pulcherrima, Erythrina glauca and Acacia berlandieri) showed significant C values above
0.96, while Arctostaphylos pungens (a temperate forest shrub) was not found to be signifi-

Fig. 2. Relationship between density of Pinus herrerae Martnez and mean annual temperature.

2013-07-09 14:47:34

288

Pablo Martinez-Antnez et al.

cantly affected by these variables, indicating

that its distribution habitat is wider.
Out of the 31 species of Quercus, seven
showed high values of C with respect to ASL,
SDAY, FFP and DD5, indicating that they
have specific requirements in these four variables; these species are: Quercus tarahumara,
Q. magnoliifolia, Q. subspathulata, Q. resinosa, Q. mcvaughii, Q. acutifolia and Q. aristata.
But only two of the 21 conifers (Pseudotsuga menziesii and Pinus arizonica)
showed absolute values of C greater than 0.90
for the variables ASL, SDAY, FFP and DD5.
This indicates that most conifers have physiological characteristics more resistant to low
temperatures than broadleaf species, and are
also more resistant to frosts.
The altitude (ASL) had a significant effect on the density of 65.28% of the species
(Appendixes II, III and IV). This behaviour
is comparable to the strong effect on the density of species shown by several of the climatic
variables. This is understandable, since ASL is
closely related to the temperature. C or t s C astel n and Isleb e (2005) and Shar ma
et al. (2009) reported also significant correlation between altitudinal gradient, micro topographical and soil conditions and the richness of tree species in India and south-eastern
Mexico. The results of this work suggest that
ASL may also be correlated to the incidence
of frosts, as indicated by the variables SDAY
and FFP.
It is worth to say that the low correlation
found between the species density and the
ASP, AST and SD as compared with climate
variables, are coincident with the results
mentioned by Mirand a and Her nnd e z
(1963) who describe the optimal conditions
of Quercus and Pinus in the temperate forests
of Mexico. Moreover, these low correlations
might also be explained by a higher tolerance
against variation of ASP, AST and SD and/or
higher intraspecific genetic differentiation
and diversity (Wehen kel et al. 2011b).
The results found in this study are consistent with those found by Ivers on and
Pras ad (1998) who reported that the abundance and distribution of 80 tree species is
regulated by several factors, such as data soil,
topography, temperature and elevation above
sea level, among others. Bud ke et al. (2006)
also reported that topography affects fertil-

journal 34.indb 288

ity and soil texture, suggesting that it also affects the abundance, frequency or density of
the plants. This work may contribute to more
specific studies to determine optimal ranges
for the abundance of trees and shrubs, either
using some multidimensional analysis, perhaps employing techniques as the ones used
by Pe ars on and D aw s on (2003), Haman n
and Wang (2006) and S ch r ag et al. (2008).
In general, climate variables appear to affect the density of broad leaf trees and shrubs
more than the density of conifers, since the
strongest correlations, close to unity, were
less frequent in the latter group of species, indicating that they have a greater plasticity to
adapt within their distribution rank, considering the variables analyzed. These results are
helpful to classify species into groups, according to the degree of correlation individually
shown with each of the variables considered
in this study, which may have practical applications.
4. CONCLUSIONS
The results of this work may be considered as a contribution to the study on stand
density and distribution of plant species of
Mexican forests. The information obtained
could also be a support to select species tolerant to certain areas of interest if their climatic
and physiographic characteristics are known,
and to support future studies for the design
of reforestation and planting programs with
better chances to succeed, either for commercial or restoration purposes. Given the global
climatic change, if the environmental variables most related to the species of interest
are known, this study may also contribute to
planning a program for assisted plant migration, by looking for similar sites for species
that currently grow in areas with specific requirements and for those which are in danger
of extinction. Overall, this work allowed detecting which of the studied species are highly
correlated with certain variables, concluding
that these species are most vulnerable to a
drastic change of those variables. However it
is still pending, for future research, conducting regression analysis to detect the groups of
variables and models that explain the density
of each species of trees and shrubs studied in
this work.

2013-07-09 14:47:34

Effect of climate and physiography on the density of tree and shrub species

AKNOWLEDGMENTS: The authors of

this paper are thankful to the National Forest
Commission, Durango delegation and to the Forest Service of the United States Department of Agriculture (FS-USDA) for their contribution with
valuable information for this study.
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Received after revision February 2013

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Effect of climate and physiography on the density of tree and shrub species

291

APPENDIX I. Species studied showing their frequency of occurrence (Freq.) and mean density in the
study area.
Species
Abies
durangensis
Martnez
Abutilon sp.

Freq. Density
(% of (Stems/
plots)
ha)

Species

Freq. Density
(% of (Stems/
plots)
ha)

0.39

0.38

Pinus cembroides
Zucc

14.63 11.62 Humb. & Bonpl.

0.11

0.18

Pinus chihuahuana
Engelm

9.04

3.87

0.54

Pinus cooperi D. Don

7.43

1.39

Pinus douglasiana
Martnez

2.44

0.43

Pinus durangensis
Martnez

26.00 27.52 Torr.

0.43

Pinus engelmannii
Carr.

17.85

0.76

Pinus herrerae
Martnez

Acacia
berlandieri
1.05
Benth
Acacia
macracantha
2.55
Humb. &
Bonpl 1
Acacia
farnesiana (L.) 1.22
Willd
Acacia
schaffneri S.
1.27
Watson
Alnus
acuminata
1.94
Kunth
Alnus
firmifolia
1.39
Fernald
Alnus
jorullensis
1.39
Kunth
Arbutus
xalapensis
40.08
Kunth2
Arctostaphylos
pungens
2.55
Kunth
Bursera
coyucensis
1.55
Bullock
Bursera
graveolens
(Kunth)
0.22
Triana &
Planch3
Bursera
simaruba (L.) 1.50
Sarg
Caesalpinia
pulcherrima
0.72
(L.) Sw
Colubrina
heteroneura
0.50
Griseb. Standl
Cupressus
lusitanica
1.83
Miller

Freq. Density
(% of (Stems/
plots)
ha)

Species
Quercus crassifolia

17.90

15.39

Quercus depressipes
Trel.

0.78

0.79

7.12

Quercus durifolia
Seemen

10.53

5.97

1.12

Quercus eduardii
Trel.

8.31

5.08

Quercus emoryi

3.33

2.39

7.04

Quercus fulva
Liebm.

5.10

2.34

6.04

4.76

Quercus gentryi
Muell.

2.05

0.55

1.39

Pinus leiophylla Schl.

22.51
& Cham.

14.28

Quercus grisea
Liebm.

15.91 23.16

0.51

Pinus lumholtzii
Robins & Ferns

18.24 13.28 Liebm.

Quercus laeta

21.34 17.66

16.28 Prez de la Rosa

Pinus luzmariae

4.27

1.74

Quercus
magnoliifolia Ne

5.71

6.11

0.64

Pinus maximinoi
H.E.Moore

0.44

0.18

Quercus mcvaughii
Spellenb.

0.61

0.39

0.47

Pinus oocarpa
Schiede

3.44

1.66

Quercus obtusata
Humb. & Bonpl.

10.14

5.18

0.24

Pinus teocote
Schlecht & Cham.

27.72 22.98 Trel.

Quercus radiata

4.93

2.69

0.41

Populus tremuloides
Michx.

0.83

0.43

Quercus resinosa
Liebm.

3.60

4.21

0.18

Prunus serotina Ehrn

0.89

0.18

Quercus rugosa Ne

9.04

5.00

0.28

Pseudotsuga
menziesii Mirb

2.88

1.55

Quercus salicifolia
Ne

2.22

1.49

1.23

Quercus acutifolia
Ne

1.22

0.39

Quercus scytophylla
Liebm.

2.88

1.66

continued overleaf

journal 34.indb 291

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Pablo Martinez-Antnez et al.

292
APPENDIX I cont.
Species

Freq. Density
(% of (Stems/
plots)
ha)

Species

Freq. Density
(% of (Stems/
plots)
ha)

Species

Freq. Density
(% of (Stems/
plots)
ha)

Erythrina
Quercus aristata
Quercus sideroxyla
0.28
0.20
0.67 0.59 Humb. & Bonpl.
34.53 34.01
glauca Willd.4
Hook. & Arn.
Guazuma
Quercus arizonica
Quercus splendens
1.61 1.08 Sarg.
4.60 3.95 Ne
0.89 0.38
ulmifolia Lam.
Juniperus
Quercus candicans
Quercus
monticola
1.88
0.71
0.61 0.42 Ne
0.83 0.68
subspathulata Trel.
5
Martnez
Juniperus
Quercus castanea
Quercus tarahumara
deppeana
0.55
0.66
35.09 14.21 Ne
0.67 0.48 Spellenb.
Steud
Juniperus flacQuercus chihuahuenQuercus urbanii
4.10 2.01 sis Trel.
5.60 10.03 Trel.
0.55 0.43
cida Schldl
Pinus arizoQuercus coccolobifoQuercus viminea
9.87 14.90 lia Trel.
6.82 3.91 Trel.
4.21 1.45
nica Engelm
Pinus ayacaQuercus conzattii
Tilia mexicana
21.01 8.24 Trel.
1.39 0.95 Schldl
2.88 2.27
huite Ehrenb
* Frequency is the percentage of plots where each species was found (Maurer et al. 2003). Density was estimated
dividing the total number of stems by the 288.64 ha represented in the 1804 plots (Kronenfeld and Wang 2007).
The Species marked with superscripts are some of the misidentifications detected according with G onz le z -E li z ond o (2012a): 1Acacia macracantha (it has not been reported in Durango, it could be A. cochliacantha), 2Arbutus
xalapensis (in Durango there are at least five tree species of Arbutus with different environmental requirements, obviously it was considered as only one), 3Bursera graveolens (it has not been reported in Durango, it could be B. penicillata) 4Erythrina glauca (it could be E. flabelliformis) and 5Juniperus montcola (the one in Durango is J. blancoi).

journal 34.indb 292

2013-07-09 14:47:34

journal 34.indb 293

0.06
-0.31

-0.36

-0.55

Pinus teocote

Pinus cooperi

0.29
0.06

0.11
-0.10

-0.32

-0.32
-0.64
0.04
0.14

-0.27
-0.01

Pinus arizonica

Juniperus deppeana

Juniperus monticola

Pinus luzmariae

Pinus lumholtzii

Pinus chihuahuana

Pinus engelmannii

0.95

0.14

-0.26

0.03
0.17

0.02

0.38

0.43
0.20

0.70

0.61

-0.10

0.07

-0.54

-0.58

-0.90

-0.05

-0.47

0.02

-0.35

-0.35

0.95
-0.34

0.13

-0.85

0.17

-0.87

0.04

0.69

0.17

0.58

0.22

0.27

-0.10

-0.15

0.15

-0.13

0.56

-0.21

0.35

0.28

0.37

0.35

0.67

-0.72

0.73

0.88

0.41

GSP
0.88

0.28

0.00

-0.08

0.42

-0.31

0.39

0.29

-0.09

-0.24

0.50

-0.43

0.35

-0.31

-0.72

-0.05

-0.08

-0.72

-0.59

-0.83

-0.37

-0.46

-0.45

-0.74

-0.62

0.60

-0.92

0.76

-0.77

0.15

-0.16

-0.26

0.55

-0.41

0.37

-0.02

-0.72

-0.04

-0.52

-0.80

-0.57

-0.37

-0.40

-0.76

-0.49

0.55

-0.94

0.78

-0.70

0.06

0.19

0.05

0.28

-0.20

0.41

-0.19

-0.66

-0.07

-0.56

-0.78

-0.21

-0.55

-0.49

-0.76

-0.71

0.80

-0.90

0.57

-0.77

-0.13

0.25

0.16

-0.26

-0.12

0.44

-0.19

-0.55

-0.06

-0.50

-0.69

-0.16

-0.58

-0.51

-0.69

-0.74

0.67

-0.83

0.36

-0.75

MTCM MMIN MTWM MMAX

-0.76
-0.67
-0.84
-0.85

-0.16

-0.57

-0.79

-0.26

-0.54

-0.55

-0.76

-0.63

0.97

-0.91

0.67

-0.77

MAT
-0.82

0.35

0.52

0.45

-0.29

0.04

0.67

0.89

0.30

0.62

0.62

0.21

0.94

-0.53

0.70

0.70

0.77

0.59

0.44

-0.66

0.83

ASL
-0.70

0.37

0.47

0.48

-0.53

-0.27

0.74

0.89

0.33

0.80

0.75

0.09

0.94

-0.83

0.84

0.75

0.78

0.61

0.45

-0.84

0.89

SDAY
-0.78

-0.34

-0.49

-0.43

0.54

0.19

-0.76

-0.92

-0.38

-0.79

-0.73

-0.23

-0.96

0.82

-0.83

-0.74

-0.76

-0.72

-0.35

0.85

-0.88

FFP
0.84

-0.46

-0.51

-0.55

0.33

0.14

-0.72

-0.92

-0.33

-0.88

-0.77

-0.06

-0.96

0.66

-0.84

-0.77

-0.83

-0.60

-0.57

0.66

-0.91

DD5
0.48

0.22

0.24

0.28

0.37

0.38

0.39

0.43

0.44

0.44

0.46

0.48

0.48

0.51

0.52

0.57

0.58

0.59

0.61

0.61

0.63

C mean
0.68

*ASP: Geographical aspect, ASL: Altitude above sea level, AST: Average slope of the terrain, SD: Depth of the soil, MAT: Mean annual temperature, MAP: Mean annual precipitation, GSP: Total precipitation in the growing season (April to September), MTCM: Mean temperature in the coldest month, MMIN: Mean minimum temperature in the coldest
month, MTWM: Mean temperature in the warmest month, MMAX: Mean maximum temperature in the warmest month, SDAY: Day of the year in which it is probable to happen
the last frost in spring, FFP: Frost free period, DD5: Degree days above 5C.

-0.11

-0.05

-0.49

-0.28

-0.18

0.39

-0.34

Pinus leiophylla
-0.33

0.02
-0.08

-0.31

Pinus cembroides

Juniperus flaccida

0.06

Pseudotsuga menziesii

Abies durangensis

0.13

0.18
0.52

Pinus oocarpa

-0.07

0.24

0.72

0.25

-0.02

0.11

Pinus durangensis

0.78
-0.70

0.43
-0.02

0.14

Cupressus lusitanica

0.52

0.50

0.08

Pinus herrerae

0.26

-0.19

0.03

0.53

Pinus douglasiana

MAP
0.88
0.82

0.10

0.21

Pinus ayacahuite

SD
-0.68
-0.16

ASP
0.04

AST
0.27

SPECIES
Pinus maximinoi

APPENDIX II. Species of conifers and their covariation (C) (eq. 1) with each variable*. Highlighted in bold are the C values that were significant with P <0.025
ordered from the highest to the lowest average absolute values (C mean). C values equal to 0.9 or higher are marked with grey.

Effect of climate and physiography on the density of tree and shrub species
293

2013-07-09 14:47:35

journal 34.indb 294

MAP
0.61
0.59
0.81
0.94
0.30
0.97
0.05
0.87
0.44
0.09
0.60
-0.86
-0.11
-0.62
0.12
0.92
0.67
0.83
0.84
0.28
0.71
-0.34
-0.40
0.47
-0.44
0.15
-0.03
0.12
-0.41
-0.75
0.25

GSP
0.61
0.70
0.80
0.96
0.32
0.97
0.26
0.89
0.38
-0.07
0.65
-0.84
-0.06
-0.61
0.39
0.97
0.66
0.84
0.85
0.34
0.78
-0.43
-0.42
0.42
-0.35
0.16
-0.04
0.12
-0.40
-0.74
0.27

MAT
0.92
0.92
-0.65
0.63
0.78
0.41
0.46
0.20
-0.62
0.72
-0.82
-0.70
-0.72
0.87
-0.62
0.17
0.09
0.08
-0.36
0.37
0.23
0.44
0.36
-0.41
0.33
0.04
0.01
-0.28
0.07
-0.09
0.03

MTCM
0.92
0.92
-0.69
0.54
0.85
0.54
-0.11
0.39
-0.56
0.60
-0.81
-0.66
-0.85
0.88
-0.75
0.06
0.50
0.25
-0.40
0.15
0.24
0.56
0.22
-0.23
0.50
0.17
0.16
-0.22
-0.08
-0.10
-0.10

MMIN MTWM MMAX

0.91
0.87
0.84
0.95
0.88
0.71
-0.60
-0.55
-0.56
0.62
0.72
0.42
0.89
0.83
0.66
0.57
0.24
0.03
0.53
-0.64
-0.56
0.52
-0.08
-0.42
-0.44
-0.57
-0.59
0.68
0.64
0.61
-0.77
-0.84
-0.83
-0.66
-0.58
-0.58
-0.79
-0.86
-0.71
0.87
0.86
0.77
-0.40
-0.83
-0.53
0.14
0.21
0.24
0.33
0.36
0.15
0.42
0.13
-0.51
-0.16
-0.27
-0.26
0.09
0.49
0.63
0.27
0.24
0.32
0.63
0.35
0.04
0.43
0.37
0.24
-0.17
-0.49
-0.50
0.42
0.11
0.14
0.00
0.02
-0.17
-0.05
-0.09
-0.11
-0.21
-0.31
-0.27
-0.10
0.20
0.30
-0.03
-0.03
-0.08
-0.14
0.09
0.19

ASL
-0.83
-0.88
-0.92
-0.93
0.86
-0.97
-0.89
-0.65
0.80
0.65
0.31
-0.22
0.38
0.14
0.08
-0.43
-0.54
-0.30
-0.24
-0.50
0.45
-0.03
-0.22
0.24
-0.03
0.65
0.27
0.13
-0.05
0.03
-0.03

SDAY
-0.92
-0.94
-0.96
-0.93
0.95
-0.98
-0.97
-0.60
0.80
0.78
0.24
-0.29
0.35
0.01
-0.57
-0.39
-0.59
-0.45
-0.32
-0.66
0.19
-0.42
-0.37
0.30
-0.47
0.64
0.35
0.03
-0.13
-0.02
0.01

FFP
0.91
0.94
0.96
0.94
-0.93
0.99
0.98
0.68
-0.80
-0.75
-0.19
0.17
-0.38
-0.08
0.47
0.49
0.66
0.50
0.40
0.69
-0.20
0.39
0.33
-0.23
0.41
-0.64
-0.35
-0.02
0.09
-0.08
0.03

DD5
0.85
0.90
0.91
0.93
-0.91
0.98
0.94
0.51
-0.85
-0.77
-0.36
0.36
-0.45
-0.01
0.36
0.41
0.41
0.16
0.07
0.46
-0.52
0.22
0.27
-0.34
0.35
-0.66
-0.29
-0.12
0.11
0.05
-0.01

*ASP: Geographical aspect, ASL: Altitude above sea level, AST: Average slope of the terrain, SD: Depth of the soil, MAT: Mean annual temperature, MAP: Mean annual precipitation, GSP: Total precipitation in the growing season (April to September), MTCM: Mean temperature in the coldest month, MMIN: Mean minimum temperature in the coldest
month, MTWM: Mean temperature in the warmest month, MMAX: Mean maximum temperature in the warmest month, SDAY: Day of the year in which it is probable to happen
the last frost in spring, FFP: Frost free period, DD5: Degree days above 5C

SD
0.20
0.00
-0.16
-0.24
-0.20
-0.28
-0.61
-0.40
0.30
0.34
0.01
-0.23
-0.24
-0.19
-0.24
-0.41
-0.01
0.11
0.59
-0.03
0.01
-0.21
0.25
-0.26
0.26
-0.28
0.36
0.21
-0.04
0.05
-0.07

C mean
0.72
0.71
0.68
0.68
0.67
0.61
0.55
0.53
0.52
0.49
0.46
0.46
0.44
0.43
0.42
0.41
0.40
0.39
0.38
0.37
0.36
0.34
0.32
0.31
0.30
0.27
0.20
0.17
0.15
0.15
0.10

ASP
-0.12
0.22
-0.31
0.24
0.64
0.05
-0.09
0.64
-0.07
0.10
0.02
0.05
0.10
0.00
-0.07
0.30
0.11
0.33
0.13
0.11
0.18
0.64
0.08
-0.03
-0.21
-0.18
0.15
0.15
0.05
-0.04
-0.07

APPENDIX III. Species of Quercus and their covariation (C) (eq.1) with each variable*. Highlighted in bold are the C values that were significant with P<0.025
ordered from the highest to the lowest average absolute values (C mean). C values equal to 0.9 or higher are marked with grey.

AST
0.56
0.44
0.64
0.43
0.21
0.62
0.64
0.63
0.05
0.11
-0.01
-0.21
-0.17
-0.10
-0.41
0.53
0.48
0.55
0.41
0.41
0.70
0.04
0.53
0.26
-0.21
0.03
0.59
0.21
-0.08
-0.01
0.15

Pablo Martinez-Antnez et al.

SPECIES
Quercus tarahumara
Quercus magnoliifolia
Quercus subspathulata
Quercus resinosa
Quercus mcvaughii
Quercus acutifolia
Quercus aristata
Quercus splendens
Quercus sideroxyla
Quercus crassifolia
Quercus obtusata
Quercus chihuahuensis
Quercus durifolia
Quercus arizonica
Quercus conzattii
Quercus urbanii
Quercus gentryi
Quercus scytophylla
Quercus candicans
Quercus viminea
Quercus castanea
Quercus depressipes
Quercus salicifolia
Quercus coccolobifolia
Quercus emoryi
Quercus rugosa
Quercus radiata
Quercus laeta
Quercus eduardii
Quercus grisea
Quercus fulva

294

2013-07-09 14:47:35

journal 34.indb 295

0.15
0.07
0.04
-0.10
0.39
0.19

0.07
0.24
-0.01
0.52
0.08
-0.17

0.40

0.62

0.65

0.70

0.50

0.63

0.13

0.16

0.83

-0.02

0.15

0.40

0.56

0.43

0.73

0.23

0.59

-0.16

-0.28

Guazuma ulmifolia

Tilia mexicana

Colubrina heteroneura

Caesalpinia pulcherrima

Acacia macracantha

Bursera coyucensis

Bursera graveolens

Populus tremuloides

Erythrina glauca

Alnus firmifolia

Acacia farnesiana

Alnus acuminate

Acacia berlandieri

Alnus jorullensis

Prunus serotina

Arbutus xalapensis

Bursera simaruba

Arctostaphylos pungens

Acacia schaffneri

0.91

0.89

-0.53

-0.78

-0.37
0.19

0.76

0.40

-0.54

-0.77

0.78

0.35

-0.28

0.92

0.91
-0.22

0.95

0.95

0.90

-0.19

0.25

-0.02

0.53

-0.21

0.37

-0.27
0.93

0.85

0.95

0.39
-0.26

0.98

0.97
-0.33

-0.59

-0.50

-0.17

0.67

0.56

0.75

0.60

0.81

0.70

0.59

0.74

0.50

0.79

0.69

0.88

0.99

0.98
0.87

GSP

MAP

0.72

-0.09

0.20

-0.07

0.22

0.00

0.38

0.54

SD

-0.66
0.73

0.75

-0.21

-0.59

-0.99

-0.63

-0.08

-0.71

-0.04

-0.55

-0.90

-0.67

0.39

-0.41

0.99

0.92
-0.64

0.93

-0.66

0.92

-0.15

-0.84

0.94

0.94
-0.82

0.92

0.97

0.89

0.98

0.99

0.99

-0.92

MTCM

0.90

0.97

0.93

0.98

0.99

0.99

-0.92

MAT

0.69

-0.55

0.05

-0.55

-0.83

-0.20

0.38

-0.47

0.93

0.82

-0.75

-0.81

0.95

0.93

0.98

0.93

0.98

0.99

0.99

-0.40

MMIN

0.77

-0.71

0.00

-0.62

-0.98

-0.76

0.11

-0.65

0.91

0.83

-0.54

-0.81

0.94

0.88

0.96

0.84

0.97

0.98

0.99

-0.91

MTWM

0.64

-0.72

0.11

-0.64

-0.89

-0.72

0.12

-0.83

0.92

0.88

-0.27

-0.69

0.93

0.77

0.93

0.75

0.95

0.96

0.98

-0.95

MMAX

0.08

0.38

-0.90

0.66

-0.17

0.51

-0.96

0.59

-0.64

0.49

-0.99

0.98

-0.85

-0.95

-0.89

-0.99

-0.98

-0.95

-0.99

-0.98

ASL

-0.08

0.17

-0.92

0.64

-0.23

0.45

-0.99

0.72

-0.76

0.66

-0.97

0.99

-0.99

-0.96

-0.95

-0.98

-0.98

-0.96

-0.99

-0.97

SDAY

0.02

-0.21

0.94

-0.63

0.21

-0.40

0.99

-0.60

0.73

-0.54

0.98

-0.99

0.99

0.96

0.95

0.98

0.98

0.97

0.99

0.98

FFP

0.08

-0.13

0.89

-0.71

0.20

-0.61

0.97

-0.79

0.72

-0.70

0.99

-0.99

0.87

0.95

0.94

0.98

0.97

0.96

0.99

0.99

DD5

0.39

0.46

0.47

0.49

0.51

0.55

0.56

0.60

0.63

0.65

0.67

0.72

0.75

0.76

0.77

0.77

0.79

0.80

0.83

0.87

C mean

*ASP: Geographical aspect, ASL: Altitude above sea level, AST: Average slope of the terrain, SD: Depth of the soil, MAT: Mean annual temperature, MAP: Mean annual precipitation, GSP: Total precipitation in the growing season (April to September), MTCM: Mean temperature in the coldest month, MMIN: Mean minimum temperature in the coldest
month, MTWM: Mean temperature in the warmest month, MMAX: Mean maximum temperature in the warmest month, SDAY: Day of the year in which it is probable to happen
the last frost in spring, FFP: Frost free period, DD5: Degree days above 5C.

0.02

-0.05

0.33

0.18

-0.07

0.27

-0.10

0.78

0.85

Abutilon sp.

ASP

AST

SPECIES

APPENDIX IV. Species of other genera and their covariation (C) (eq. 1) with each variable*. Highlighted in bold are the C values that were significant with
P<0.025 ordered from the highest to the lowest average absolute values (C mean). C values equal to 0.9 or higher are marked with grey.

Effect of climate and physiography on the density of tree and shrub species
295

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