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POLISH JOURNAL OF ECOLOGY

(Pol. J. Ecol.)

61

2

283–295

2013

Regular research paper

Pablo MARTÍNEZ-ANTÚNEZ 1, Christian WEHENKEL 2, J. Ciro HERNÁNDEZ-DÍAZ 2 *,
Martha GONZÁLEZ-ELIZONDO 3, J. Javier CORRAL-RIVAS 4, Alfredo PINEDO-ÁLVAREZ 2
Doctorado Institucional en Ciencias Agropecuarias y Forestales, Universidad Juárez del Estado de
Durango, Km 5.5 Carretera Mazatlán, 34120 Durango, México
2
Instituto de Silvicultura e Industria de la Madera, Universidad Juárez del Estado de Durango, Km 5.5
Carretera Mazatlán, 34120 Durango, México; *e-mail: jciroh@ujed.mx (corresponding author)
3
Instituto Politecnico National, CIIDIR, Unidad Durango. Sigma 119 Col. 20 de Noviembre II.
Durango, 34220 México
4
Facultad de Ciencias Forestales, Universidad Juárez del Estado de Durango, Río Papaloapan y Blvd.
Durango s/n, Col. Valle del Sur, 34120 Durango, México
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 á 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
(Koż uchowsk i and Teg i r mend ž i ć 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

2013-07-09 14:47:33

2011a. 2005. 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. G op araju and Jha 2010). small portions of temperate mesophytic forest. so. Density of a species is defined as the number of individuals per unit area (Kronenfeld and Wang 2007). In the Sierra Madre Occidental mountain range that crosses the State there are woodlands of Pinus. The frequency of occurrence of a species has been measured as the percentage of incidence of the species in a given number of sites. For example. the data of the INFyS is mainly reported at the level of conglomerate. Pachau r i and Ja l low 2007. 40% of the State has a dry and semi-dry climate. the third to the southeast direction Az 120° and the fourth to the southwest Az 240°. threatening the survival of ecosystems and species (IPCC-WGI 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. to perform the National Inventory of Forests and Soils (INFyS 2004−2009). 2010. especially where Pinus is the dominant genus.284 Pablo Martinez-Antúnez et al. mean temperature in the warmest or the coldest month. 2013-07-09 14:47:33 . (2003) found that more frequent species are also more abundant. distribution and density of species. G op araju and Jha 2010). known as ‘sites’. 2008. Durango is located to the northwest of Mexico between 26°53’ and 22°16’ North and between 102°29’ and 107°16’ West (Fig. Several authors agree that the increase in global temperature affect biodiversity at different scales and ways. Recent research suggests that there has been a change in the global climate system. C onde et al. in some areas forest fires occur in periods of drought (Wehen kel et al. 2011). 2011). 14% very dry. This knowledge may be helpful to define eco-physiographic areas useful to journal 34. 2004. in forest regions forestry is a major economic activity for many people (Návar 2004).indb 284 face a possible drastic environmental change and to strengthen other technical or scientific purposes (Tcheb a kov a et al. PinusQuercus. while in non-forest areas many shrub and tree species are important for medicinal purposes. which has an approximate area of 123. Quercus-Pinus. Vegetation studies have been conducted from several approaches. in the remaining of this report the word ‘plot’ is utilized instead of conglomerate. 2011). 2007). the second to the north at Az 0°. including: diversity. geographically shifting plant and animal communities and altering the ecological niche of organisms and the ecosystem function (Desai et al. MATERIALS AND METHODS The study was conducted in the state of Durango. Peters on and Vieg l ais 2001. 2012b). Zhu et al. also have economic and social importance. 2008. are subject to timber harvesting and.317 km2. 2006). Sáenz-R omero et al. and date of last spring frost. Z hu et al. commercial or other scientific purposes (G on zá le z-E l izondo et al. The study explores relationships between tree species density to climatic variables such as accumulated precipitation in the growing season. and every 20 km in arid and semiarid regions (CONAFOR 2009). extinction (Hug hes 2000.14 m. G onzá le z-E lizondo et al. 1). 11% sub humid-warm and only 1% humid-temperate (INEGI 2012). 2. Some of these studies document changes in environmental conditions over short periods of time. besides fulfilling ecological functions such as habitat for wildlife reproduction and feeding. frequency. In the region of the valleys there are mainly desert and semi-desert zones. The sites were distributed into the conglomerate with an equidistance of 45. which are seldom used for inferring presence of plant species. the first site was located in the centre of the conglomerate. It has been stated that trees and shrubs. 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. However. The 1804 primary sampling units (conglomerates) utilized in this study were established in most of the State by the National Forest Commission (CONAFOR 2009). Ait ke n et al. Maurer et al. and on the western flanks some tropical deciduous and semi deciduous forests (G onzá le z-E lizondo et al. protection against soil erosion. Many of the forests in the study area. retention of water resources. 34% sub humid-temperate.

2008. A total of 329 species of trees and shrubs were found in the 1804 studied plots.edu/climate/) which models climate data from 4000 weather stations of Mexico.99 MAT °C 3. south.9 MTCM °C 1.R omero et al. Descriptive statistics of the independent variables*. mean minimum temperature in the coldest month (January) (MMIN. mean maximum temperature in the warmest month (June) (MMAX. east. Thus.5 cm at the height of 1.04% (since it was found in 108 out of the 1804 plots) and a mean density of 4.47 3. north-east. 2010).5 11. MAT: Mean annual temperature.47 MTWM °C 6. north-east. Following Krone n feld and Wang (2007). mean temperature in the warmest month (June) (MTWM.9 26. journal 34. east. MAP: Mean annual precipitation. ASP: Geographical aspect (zenith.85 MAP mm 250 1444 878 251. m). average length Table 1.52 AST % 0 98 25 16.8 40. GSP: Total precipitation in the growing season (April to September). Four physiographic and site specific variables considered in this work include: the average slope of the terrain (AST.3 m above the ground level were counted and identified at the species level. all the stems of trees and shrubs. Pinus herrerae presented a frequency of 6. °C). MMIN: Mean minimum temperature in the coldest month.61 SDAY days 0 196 109.7 18. South of the United States of America. The climate variables obtained from that source for each of the 1804 INFyS plots. Over 80% of these species were found in the plots located on the Sierra Madre Occidental mountain range.1 14 3.82 ASL m 206 4502 2219 512. altitude 285 above sea level (ASL. moscowfsl. south-east. AST: Average slope of the terrain. Belize and Cuba.5 -0.1 26. the final data set was composed of 72 species of trees and shrubs (Appendix I).D. showing the frequency of occurrence (Maurer et al. MTCM: Mean temperature in the coldest month. 10 climate variables considered important according to the algorithms of R e h fel dt (2006). DD5: Degree days above 5°C. north-west and south-west) and the soil depth (SD. °C). west. 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. total precipitation in the growing season (April to September (GSP. °C).91 FFP days 43 365 196. north. were: mean annual temperature (MAT. with records from 1976 to 1990 (Cro ok ston et al. mean annual precipitation (MAP. following the criterion of finding at least 50 individuals of each species in the 1804 plots. south. mm). 13 species were later discarded because their taxonomic identification was very doubtful. south-east. In addition. in cm) (CONAFOR 2009). MTWM: Mean temperature in the warmest month. FFP: Frost free period.3 30. dominant aspect or geographic orientation of the ground (ASP: zenith. west.89 44.2 21. developed by Hutch i ns on (2004).53 3.49 DD5 °C 16 253 109. °C).84 GSP mm 190 1068 667 167. S áen z .Effect of climate and physiography on the density of tree and shrub species In each plot. northwest and south-west). ASL: Altitude above sea level. Nevertheless.76 stems per hectare. mean temperature in the coldest month (MTCM.91 MMAX °C 12.48 SD cm 0 110 38 23. from which 85 were selected for further analyses. mm).D.35 * S. These data are modelled using the ANUSPLIN software.07 48. Variable Units Minimum Maximum Mean S.9 3.: Standard deviation. Guatemala. ASP * 0 315 128. whose diameter was greater than or equal to 7. °C). SD: Depth of the soil.67 4.27 109.06 75.indb 285 2013-07-09 14:47:33 . 2003) and the mean density of each species per hectare in the study area. were obtained from the website of the Forest Service of the Department of Agriculture in the United States (http://forest.94 MMIN °C -7. north. The whole list of studied species is presented in Appendix I. For example. the number of stems recorded in each plot is treated in this paper as density.2 8.wsu. in %).

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. 2006). and Tilia mexicana presented the highest correlations simultaneously with several variables and also. days). ranges between −1 and +1. By definition. 2005.65 each of these three variables.66. East 90. Formally: Fig. The denominator of Eq. Table 1 provides a summary of the conditions prevailing in the study area. The covariation C journal 34. C is undefined (Gre gor ius et al. in terms of the basic values of the variables.286 Pablo Martinez-Antúnez et al. (1) is the summation of absolute values of the indicated products. To evaluate the correlation between the density of species and ASP (which is not a numerical variable) using the same method. Analyzing by species group. °C). R ehfeldt et al. 2007). MTWM. 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. The species Abutilon sp. (Yi-Yj) = difference in density value of the variable Y. between the i-th and j-th plots. North 1. RESULTS AND DISCUSSION The estimated covariation values of Gregorius (C) showed that 88% of the studied species have a significant correlation (P <0. between the i-th and the j-th plots. Some species of other genera different than Quercus or conifers (Appendix 2013-07-09 14:47:33 . and MTCM.71. conifers (Appendix II) and Quercus (Appendix III) showed little correlation with AST. If its denominator is zero. 3. The climatic variables that apparently largely affect the density of most of the studied species are: SDAY with an average C = 0. West 270. Given that there was not found linearity or a normal distribution of the data. indicating that they may equally be found in varying degrees of slope.71. FFP with C = 0.69. South East 135. (2007). each aspect was assigned a numerical value indicating the predominant azimuth: Zenithal 0. the largest mean values of C (mean C). North East 45. Location of the study area. 1. Julian date of the last frost spring (SDAY.025) with at least five variables of climate and physiography (Appendixes II. ASL with C = 0. of frost-free period (FFP.indb 286 (1) where: C = Covariation between each species X and variable Y. MAT with average values of C = 0. Guazuma ulmifolia. All of these variables are considered critical to the full development of plants (Tcheb a kova et al. In order to test the probability that the observed degrees of covariation C are produced solely by random events rather than directed forces. and North West 315. There thus exists a strictly monotonic relationship (but not necessarily linear) between the two variables. days) and degree-days above 5°C (DD5. III and IV). South 180. two ordinal variables X and Y show entire covariation if one variable consistently increases or consistently decreases as the other variable increases.. where C = 1 corresponds to an entirely positive covariation and C = −1 to a strictly negative covariation. South West 225. (Xi-Xj) = difference in climate values of the specie X. DD5 with C = 0.

the stronger effect of ASL. implying that their respective habitats are smaller. Out of the 21 species of conifers only one (Abies durangensis). Relationship between density of Pinus herrerae Martínez and mean annual temperature. SDAY. 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. On the other hand it is observed that MAT shows significant correlations with 71. Only seven of them showed significant correlation with this variable. FFP and DD5 seem to have the same pattern of behaviour. while Arctostaphylos pungens (a temperate forest shrub) was not found to be signifi- Fig. Also in regard to SD. SDAY. which was also observed by Silva-Flores and Wehenkel (unpublished). it was decided to report about all of them. and two trees (Tilia mexican and Bursera coyucensis ) of tropical forest.4% of the group of conifers (Appendix II). where five tropical forest shrubs (Abutilon sp. Colubrina heteroneura. since one of the contributions of this work is that. Out of them.4% for Quercus (Q. showing C values between 0. 2013-07-09 14:47:34 . In the same order. As happened with the variables that directly indicate temperature (MAT. Tilia mexicana and Colubrina heteroneura. MTCM. acuminata and A. three are of the genus Quercus. they include: two shrubs (Colubrina heteroneura and Caesalpinia pulcherrima). among the other genres (Appendix IV) the effect of precipitation was more remarkable.) showed C of 0. the percentages of absolute values of C equal to or greater than 0. considering the absolute value of C in most of the analyzed species. however.Effect of climate and physiography on the density of tree and shrub species IV) were significantly (P <0. Erythrina glauca and Acacia berlandieri) and four of the 14 trees (Guazuma ulmifolia.025) correlated with AST. MTWM and MMAX) and those related to precipitation (MAP and GSP) again.90 were respectively 9. MTWM and MMAX. Q. Two other tropical shrubs (Abutilon glauca and Erythrina sp. and only three out of the 31 oaks (Quercus resinosa.96.73.62 and 0. The above is corroborated by noting that similar correlations to those presented by the three groups of species for MAT. The ASP does not seem to be a limiting factor for the habitat of most species. urbanii) reached C values higher than 0. Juniperus monticola being the only one showing a significant high C value (0. Caesalpinia pulcherrima.. MMIN. MMIN. were also observed with respect to MTCM. A.85 respectively with AST. which indicates that they have specific requirements of precipitation.90. magnoliifolia) and 55. FFP and DD5 was found in the species listed in Appendix IV. 2.90). and a temperate forest tree (Prunus serotina). in Appendixes II. Variables ASL. journal 34..indb 287 287 Pines and oaks showed less requirements with respect to precipitation that with regard to temperature. acutifolia and Q. 6. Alnus jorullensis. But. Erythrina glauca and Acacia berlandieri) showed significant C values above 0.0% for other genres (Guazuma ulmifolia. It was found that some of these variables are inter-correlated. since it is the variable less correlated with the density of the 72 species analyzed.83 and 0. it was found no evidence of a large effect on the density of the studied species. which is distributed entirely in the mountainous region of the State. which means that the latter group is more demanding in terms of the specific temperature requirements for their existence. firmifolia) showed values of C greater than 0.5% for conifers (Cupressus lusitanica and Pinus herrerae). since three of the six shrubs (Abutilon sp. Some correlations were negative and others positive.0% of the group of other genera (Appendix IV). 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. with 51.6% of the group of Quercus (Appendix III) and with 85. but these values were not significant. tarahumara and Q. among others).90 with respect to these two variables.

FFP and DD5. suggesting that it also affects the abundance. this study may also contribute to planning a program for assisted plant migration. acutifolia and Q. cantly affected by these variables. Out of the 31 species of Quercus. as indicated by the variables SDAY and FFP. topography.28% of the species (Appendixes II. either using some multidimensional analysis. perhaps employing techniques as the ones used by Pe ars on and D aw s on (2003). 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. and are also more resistant to frosts. AST and SD and/or higher intraspecific genetic differentiation and diversity (Wehen kel et al. these low correlations might also be explained by a higher tolerance against variation of ASP. among others. 2011b). FFP and DD5. close to unity. 4. aristata. AST and SD as compared with climate variables. since ASL is closely related to the temperature. were less frequent in the latter group of species. Overall. indicating that its distribution habitat is wider. concluding that these species are most vulnerable to a drastic change of those variables. The altitude (ASL) had a significant effect on the density of 65. 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. climate variables appear to affect the density of broad leaf trees and shrubs more than the density of conifers. These results are helpful to classify species into groups. SDAY. C or té s C astel án and Isleb e (2005) and Shar ma et al. (2006) also reported that topography affects fertil- journal 34. However it is still pending. Bud ke et al. these species are: Quercus tarahumara. Q. indicating that they have a greater plasticity to adapt within their distribution rank.288 Pablo Martinez-Antúnez et al. subspathulata.90 for the variables ASL. Q. indicating that they have specific requirements in these four variables. if the environmental variables most related to the species of interest are known. Haman n and Wang (2006) and S ch r ag et al. micro topographical and soil conditions and the richness of tree species in India and south-eastern Mexico. and to support future studies for the design of reforestation and planting programs with better chances to succeed.indb 288 ity and soil texture. But only two of the 21 conifers (Pseudotsuga menziesii and Pinus arizonica) showed absolute values of C greater than 0. Moreover. It is worth to say that the low correlation found between the species density and the ASP. resinosa. which may have practical applications. (2008). (2009) reported also significant correlation between altitudinal gradient. Q. are coincident with the results mentioned by Mirand a and Her nánd e z (1963) who describe the optimal conditions of Quercus and Pinus in the temperate forests of Mexico. according to the degree of correlation individually shown with each of the variables considered in this study. for future research. by looking for similar sites for species that currently grow in areas with specific requirements and for those which are in danger of extinction. The results of this work suggest that ASL may also be correlated to the incidence of frosts. seven showed high values of C with respect to ASL. This is understandable. since the strongest correlations. either for commercial or restoration purposes. III and IV). 2013-07-09 14:47:34 . This behaviour is comparable to the strong effect on the density of species shown by several of the climatic variables. This work may contribute to more specific studies to determine optimal ranges for the abundance of trees and shrubs. SDAY. this work allowed detecting which of the studied species are highly correlated with certain variables. mcvaughii. temperature and elevation above sea level. such as data soil. In general. considering the variables analyzed. Q. frequency or density of the plants. 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. This indicates that most conifers have physiological characteristics more resistant to low temperatures than broadleaf species. magnoliifolia. 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. Q. Given the global climatic change.

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5.39 Fernald Alnus jorullensis 1.04 Quercus fulva Liebm.33 2.54 Pinus cooperi D.indb 291 2013-07-09 14:47:34 . Density (% of (Stems/ plots) ha) Species Freq.21 0.24 Pinus teocote Schlecht & Cham. 1. 2.94 Kunth Alnus firmifolia 1.88 1.05 0.50 Sarg Caesalpinia pulcherrima 0.39 0.Moore 0.38 Pinus cembroides Zucc 14.66 Quercus obtusata Humb.51 Pinus lumholtzii Robins & Ferns 18.04 4.22 1.93 2.85 0.27 1.05 Benth Acacia macracantha 2.74 Quercus magnoliifolia Née 5. 0. 22.43 Pinus durangensis Martínez 26.89 0.55 Humb.87 0.39 Pinus leiophylla Schl.39 Quercus depressipes Trel.53 5.63 11.39 7. 14.39 Quercus scytophylla Liebm.88 1.61 0.52 Torr. 0. Density (% of (Stems/ plots) ha) 0.44 0.44 1. Species studied showing their frequency of occurrence (Freq.27 Watson Alnus acuminata 1.64 Pinus maximinoi H.18 Pinus chihuahuana Engelm 9.08 Quercus emoryi 3.55 1.08 Kunth2 Arctostaphylos pungens 2. 10.14 5.12 Quercus eduardii Trel.66 continued overleaf journal 34.31 5.66 16.49 1.23 Quercus acutifolia Née 1. 0.E. 0.00 27.43 Quercus resinosa Liebm.) Sw Colubrina heteroneura 0.10 2.97 1.43 1. & Bonpl 1 Acacia farnesiana (L.) and mean density in the study area. 27.11 0. 17.78 0.04 5.22 Willd Acacia schaffneri S. 2.98 Trel.18 Quercus mcvaughii Spellenb.83 0.76 Quercus gentryi Muell.Effect of climate and physiography on the density of tree and shrub species 291 APPENDIX I.55 Kunth Bursera coyucensis 1.47 Pinus oocarpa Schiede 3.18 0.28 Pseudotsuga menziesii Mirb 2.55 Quercus salicifolia Née 2.60 4.) 1.34 17.00 0. Freq.62 Humb.44 0.76 Pinus herrerae Martínez Acacia berlandieri 1.91 23.28 Pérez de la Rosa Pinus luzmariae 4. Standl Cupressus lusitanica 1.28 Liebm.50 Griseb.12 Quercus durifolia Seemen 10.24 13.28 Quercus grisea Liebm.39 Kunth Arbutus xalapensis 40.39 Pinus douglasiana Martínez 2.22 0.79 7. 15. Species Abies durangensis Martínez Abutilon sp.72 22.41 Populus tremuloides Michx.18 Quercus rugosa Née 9.16 0.72 (L.) 1.51 & Cham. Quercus radiata 4.18 Prunus serotina Ehrn 0. 8.90 15.55 Bullock Bursera graveolens (Kunth) 0.11 0. & Bonpl. Density (% of (Stems/ plots) ha) Species Quercus crassifolia 17.69 0.04 3. & Bonpl.43 Pinus engelmannii Carr. Quercus laeta 21.71 6.83 Miller Freq.22 Triana & Planch3 Bursera simaruba (L.39 0. 3. 0. Don 7.34 6.

60 10. Density (% of (Stems/ plots) ha) Species Freq.10 2. 5 Martínez Juniperus Quercus castanea Quercus tarahumara deppeana 0.03 Trel.60 3. 1.67 0. Density (% of (Stems/ plots) ha) Species Freq.01 glauca Willd.91 Trel.01 8. Species Freq. Guazuma Quercus arizonica Quercus splendens 1. penicillata) 4Erythrina glauca (it could be E.indb 292 2013-07-09 14:47:34 .21 Née 0.59 Humb. obviously it was considered as only one). journal 34.66 35. cochliacantha).53 34.21 1. 0.01 sis Trel.45 nica Engelm Pinus ayacaQuercus conzattii Tilia mexicana 21.83 0. 3Bursera graveolens (it has not been reported in Durango.38 ulmifolia Lam.55 0.24 Trel.64 ha represented in the 1804 plots (Kronenfeld and Wang 2007). 2003).61 1. 2Arbutus xalapensis (in Durango there are at least five tree species of Arbutus with different environmental requirements. Steud Juniperus flacQuercus chihuahuenQuercus urbanii 4. 6.42 Née 0.Pablo Martinez-Antúnez et al.88 2.88 0.87 14. 4.27 huite Ehrenb * Frequency is the percentage of plots where each species was found (Maurer et al.95 Schldl 2. blancoi).20 0.95 Née 0. flabelliformis) and 5Juniperus montícola (the one in Durango is J. The Species marked with superscripts are some of the misidentifications detected according with G onzá le z -E li z ond o (2012a): 1Acacia macracantha (it has not been reported in Durango.55 0.4 Hook.39 0.89 0.90 lia Trel. Juniperus Quercus candicans Quercus monticola 1.67 0.68 subspathulata Trel.43 cida Schldl Pinus arizoQuercus coccolobifoQuercus viminea 9. 292 APPENDIX I – cont. 4. Density (% of (Stems/ plots) ha) Erythrina Quercus aristata Quercus sideroxyla 0. & Arn.71 0. it could be B. Density was estimated dividing the total number of stems by the 288.08 Sarg. it could be A. & Bonpl.48 Spellenb.82 3. 34.09 14.61 0. 5.28 0.

22 0.10 0.61 0.12 0.34 -0.63 0.26 -0.44 0.78 -0.52 0.74 0.14 -0.77 0.70 0.88 FFP 0.94 -0.01 Pinus arizonica Juniperus deppeana Juniperus monticola Pinus luzmariae Pinus lumholtzii Pinus chihuahuana Pinus engelmannii 0.67 -0.72 -0.77 MAT -0.57 -0.55 -0.72 -0.95 0.70 0.85 0.96 0.90 0.59 -0.35 0.44 -0.02 0.80 -0. 1) with each variable*.52 Pinus oocarpa -0.19 0.47 0.25 -0.79 -0. C values equal to 0. Highlighted in bold are the C values that were significant with P <0.04 AST 0.09 0.57 0.58 0.55 -0.53 Pinus douglasiana MAP 0.06 -0.66 -0.08 Pinus herrerae 0.89 SDAY -0.89 0.92 0.84 0.43 0.72 -0.00 -0.55 Pinus teocote Pinus cooperi 0.48 -0.32 -0.29 -0.49 -0.51 0.62 0.74 -0.22 0.54 -0.55 -0.45 -0.06 -0.17 -0.96 0.43 0.28 0.15 -0.49 -0.24 0.20 0.33 -0.63 C mean 0.02 0.50 -0.67 -0.70 0.24 0.67 -0.35 -0.57 0.83 -0. GSP: Total precipitation in the growing season (April to September).75 0.48 0.35 0.10 0.72 -0.76 -0.60 -0.83 ASL -0.10 -0.05 -0.53 0.83 0.04 -0.92 -0.89 0.59 0.31 -0.18 0.78 0.20 0. AST: Average slope of the terrain.80 0.28 0. DD5: Degree days above 5°C.70 0.49 0.51 -0.16 -0.03 0.28 -0.15 0.61 0.54 -0.13 0.48 0.27 -0.77 0.07 0.64 0.62 0. -0.55 -0.11 -0.08 -0.41 -0.92 -0.51 -0.56 -0.82 0.41 0.85 -0.94 0.04 0.21 0.80 -0.55 -0.indb 293 0.55 0.66 0.36 -0.39 0.34 0.69 -0.73 -0.39 -0.35 0.32 -0.37 -0.38 0.25 0.57 -0. FFP: Frost free period.88 0.43 -0.61 0.57 -0. Species of conifers and their covariation (C) (eq.28 0.76 -0.10 -0.83 0.02 -0.88 -0.75 MTCM MMIN MTWM MMAX -0.70 0.journal 34. MTCM: Mean temperature in the coldest month.37 0. MMAX: Mean maximum temperature in the warmest month.025 ordered from the highest to the lowest average absolute values (C mean).38 0.05 -0.16 -0.66 -0. MAP: Mean annual precipitation.69 -0. SDAY: Day of the year in which it is probable to happen the last frost in spring. MTWM: Mean temperature in the warmest month.84 0.03 0.26 0.05 -0.50 -0.17 0.27 SPECIES Pinus maximinoi APPENDIX II.54 0.29 0.59 0.44 -0.43 0.08 -0.84 -0.82 -0.43 0.21 0.76 -0.27 0.72 -0. ASL: Altitude above sea level.14 -0.26 0.76 -0.37 -0.77 -0.16 ASP 0.76 -0.42 -0.77 -0.19 -0.70 0.85 -0.13 -0.95 -0.19 -0.45 -0.58 -0.16 -0.35 0.35 -0.30 0.26 -0.16 -0.27 -0. MMIN: Mean minimum temperature in the coldest month.19 0.78 -0. SD: Depth of the soil.33 0.09 -0.46 0.72 0.67 -0.26 -0.40 -0.76 -0.69 0.91 DD5 0.19 -0.77 -0.58 -0.41 GSP 0.13 0.06 Pseudotsuga menziesii Abies durangensis 0.82 0.56 -0. MAT: Mean annual temperature.88 0.37 0.29 0.45 -0.79 -0.08 0.52 0.39 0.06 0.68 *ASP: Geographical aspect.83 -0.14 -0.31 0.83 -0.02 -0.04 0.49 -0.66 -0.33 0.74 -0.37 -0.38 -0.88 0.74 0.33 0.06 -0.58 0.68 -0.97 -0.46 -0.9 or higher are marked with grey.71 0.17 0.35 0.14 Cupressus lusitanica 0.73 0.21 Pinus ayacahuite SD -0.31 -0.48 0.50 0.31 Pinus cembroides Juniperus flaccida 0.37 0.91 0.24 0.94 -0.15 -0.11 -0.62 0.23 -0.60 -0.13 0.72 0.47 0.52 -0.04 0.84 -0.46 -0.05 0.18 0.07 -0.67 0.53 -0.84 -0.02 -0.36 -0.44 0.07 -0.75 0.11 Pinus durangensis 0.61 -0.90 -0. Effect of climate and physiography on the density of tree and shrub species 293 2013-07-09 14:47:35 .87 0.21 -0.78 -0.06 0.78 -0.76 -0.02 0.34 Pinus leiophylla -0.28 -0.52 0.

02 0.63 0.65 0.04 -0.15 0.15 0.03 -0.50 0.40 0.36 0.66 -0.86 0.80 0.83 0.92 -0.86 -0.38 0.30 -0.70 -0.42 0.15 0.16 -0.40 0.23 -0.42 0.86 -0.21 0.20 -0.17 -0.journal 34.22 0.08 -0.01 FFP 0.00 -0.08 -0.36 0.38 -0.21 0.30 0.05 -0.31 -0.45 -0.27 0.31 -0.85 0.02 0.09 -0.36 0.86 -0.32 -0.34 -0.95 0.24 0.64 -0.81 0.38 0.65 0. 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 .32 0.33 0.11 0.39 0.20 0.46 0.20 0.03 0.61 0.46 -0.13 -0.24 -0.97 -0.05 0.69 -0.09 0.01 -0.35 0.72 0.07 0.42 0.61 0.00 -0. Highlighted in bold are the C values that were significant with P<0.59 -0.12 0.01 0.12 -0.37 0.72 0.84 0.53 0.49 -0.18 0.35 0.98 0.02 0.65 -0.05 -0.12 0.95 -0.09 -0.16 -0.08 0.37 0.9 or higher are marked with grey.60 -0.21 -0.09 0.56 0.98 0.24 -0.36 -0.06 0.44 0.60 -0.64 0.41 0.92 -0.52 -0.24 0.83 -0.26 -0.11 0.26 -0.82 -0.44 0.24 -0.77 -0.58 -0. SD: Depth of the soil.91 0.05 -0.59 -0.92 -0.54 -0.41 0.39 -0. C values equal to 0.93 0.11 -0.10 ASP -0.94 -0.18 0.63 0.13 -0.03 -0.43 0.12 0.04 0.26 0.58 -0.26 0.15 0.54 -0.66 0. MTWM: Mean temperature in the warmest month.44 0.10 MMIN MTWM MMAX 0.01 -0.27 MAT 0.78 -0.16 -0. DD5: Degree days above 5°C SD 0.64 0.17 0.08 -0.21 -0.25 -0.60 -0.51 -0.56 0. SDAY: Day of the year in which it is probable to happen the last frost in spring.80 0.93 -0.42 -0.30 -0.98 -0. FFP: Frost free period.62 0.1) with each variable*.41 0.66 0.75 0.22 0.06 -0.08 0.97 0.61 0.25 GSP 0.47 -0.88 -0.22 -0.22 0. ASL: Altitude above sea level.03 -0.36 0.33 0.35 0.12 -0.24 -0.46 0.30 0.24 -0.79 -0.62 0.07 0.70 0.03 0.21 0.24 0.64 -0.84 -0.10 0.44 -0.91 0. Species of Quercus and their covariation (C) (eq.20 0.05 0.43 -0.87 -0.03 MTCM 0.05 0.27 -0.36 0.72 0.66 0.93 0.68 0.57 -0.08 -0.21 -0.71 -0.04 0.25 -0.11 -0.45 -0.35 0.22 -0.75 0.43 0.24 -0.89 0.23 0.10 0.47 0.03 SDAY -0.02 -0.11 0.48 0.09 0.32 0.87 0.17 -0.00 0.42 0.97 0.16 0.52 0.85 0.46 0.89 -0.66 0.13 -0. AST: Average slope of the terrain.33 0.80 0.44 0.26 0.94 -0.57 -0.21 0.41 0.53 0.56 0.34 0.19 -0.37 0.09 0.56 0.43 0.24 0.78 0.84 -0.53 -0.71 -0.30 0.08 -0.13 0.42 -0.64 0.34 0.85 0. MAP: Mean annual precipitation.11 -0.81 -0.20 0.025 ordered from the highest to the lowest average absolute values (C mean).87 0.27 0.36 0.10 -0.17 -0.96 0.15 0.83 0.94 0.60 0.93 0.23 0.64 0.01 0. MTCM: Mean temperature in the coldest month.05 0.01 *ASP: Geographical aspect.83 -0.44 0.92 -0.66 -0.23 0.39 0.10 0.53 0.16 -0.39 0.15 -0.17 0.19 -0.43 -0.41 -0.49 0.05 -0.63 0.30 0.88 -0.10 -0.92 0.99 0.05 -0.01 0. GSP: Total precipitation in the growing season (April to September).14 0.61 0.67 0.53 0.90 0.08 -0.97 -0.71 0.41 -0.72 -0.62 0.33 -0.49 0.09 0.75 -0.83 -0.56 0.27 -0.38 -0.50 0.07 0.17 -0.03 0.77 -0.03 DD5 0.88 0.34 0.62 0.27 -0. MMIN: Mean minimum temperature in the coldest month.55 0.41 0.57 0.55 -0.77 -0.45 -0.20 -0.11 0.91 0.07 C mean 0.70 0.47 0.24 -0.69 0.59 0.65 0.19 ASL -0.92 0.40 0.40 0.04 0. MAT: Mean annual temperature.92 0.28 0.61 -0.41 0.31 0.01 -0.28 -0.08 -0.01 -0.29 0.41 -0.74 0.04 0.71 0. MMAX: Mean maximum temperature in the warmest month.32 0.19 0.14 0.22 -0.49 0.40 -0.28 0.37 0.84 0.01 -0.50 0.64 0.35 -0.34 0.94 0.04 0.41 0.03 -0.42 0.65 0.40 0.87 0.39 -0.59 0.50 0.35 -0.85 0.28 0.68 0.40 -0.21 -0.64 -0.03 0.29 -0.63 0.31 0.indb 294 MAP 0.09 0.80 -0.52 0.96 -0.59 0.14 0.03 -0.66 -0.15 0.07 -0.17 0.03 -0.36 -0.03 0.21 -0.07 0.41 0.78 0.55 0.62 0.50 0.51 -0.84 0.96 0.27 0.61 -0.64 0.67 0.89 0.15 Pablo Martinez-Antúnez et al.14 0.97 0.30 -0.11 0.94 0.68 -0. AST 0.54 0.24 0.07 APPENDIX III.11 0.85 -0.42 -0.68 0.91 0.

99 -0.08 0.75 0.97 0.92 -0. C values equal to 0.64 0.93 0.59 -0.94 0.50 -0.46 0.65 0.55 -0.98 -0.17 0.98 ASL -0.96 0.60 0.95 -0.89 -0.07 0. FFP: Frost free period.88 0.92 MAT 0.99 DD5 0. MAP: Mean annual precipitation.94 -0.02 0.93 0.63 0.77 -0.72 0.02 0.98 -0.75 -0.82 -0.70 0.88 -0.99 -0.17 -0.98 0.79 0.20 -0.20 -0.02 -0.89 -0.28 0.49 -0.33 -0.63 0.54 0.76 0.83 -0.04 -0. ASL: Altitude above sea level.71 -0.70 0. MAT: Mean annual temperature.95 0.61 0.39 -0.83 0.99 -0.11 -0.journal 34.97 -0.18 -0.66 0.71 0.79 0.62 0. 0.99 0.92 -0.96 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 0.72 -0.26 0. ASP AST SPECIES APPENDIX IV.55 0.59 -0.78 0.38 -0.67 0. 1) with each variable*.98 0.99 -0.99 -0.17 0.73 0.64 0.99 -0.98 FFP 0. Highlighted in bold are the C values that were significant with P<0.66 0.98 0.96 -0.72 0.19 0.94 0.22 0.99 -0.90 -0.99 -0.98 -0.95 0.82 0.10 0.90 -0.72 0.15 -0.11 -0.97 -0.40 MMIN 0.99 -0.75 0.74 0.13 0.47 0.92 0.94 -0.87 C mean *ASP: Geographical aspect.41 0.89 0.25 -0.35 -0.99 0.27 -0.04 -0.81 0.08 0.55 0.15 0.54 SD -0.59 -0.59 0.52 0.99 0.72 -0.49 0.27 -0.39 0.99 0.97 0.10 0.21 -0.40 0.93 0.39 0.07 0.85 -0.17 0.99 -0.93 0.99 0. DD5: Degree days above 5°C.54 -0.83 0.23 0.64 -0.77 0.99 0.53 -0.98 0.73 0.98 -0.43 0.96 0. MMIN: Mean minimum temperature in the coldest month.93 0.70 0. SD: Depth of the soil.99 0.83 -0.91 -0.00 -0.99 -0.66 -0.98 0.77 0.87 0.40 0.92 0.97 0.08 -0.40 0.05 -0.69 0.59 -0.96 0.92 MTCM 0.21 0.88 0.13 0.65 0. MTCM: Mean temperature in the coldest month.51 -0.95 0.76 0.76 0.98 0.33 0.81 0.64 -0.56 0.81 0.95 -0.83 -0.65 0.67 0.77 0.97 0.23 0.91 0.45 -0.90 0.98 0.27 0.56 0.97 0.96 0.69 0.93 -0.76 0.99 0.99 0.95 0.64 -0.50 0.01 0.75 0.73 -0.72 -0. MMAX: Mean maximum temperature in the warmest month.97 SDAY 0.63 -0.55 -0.60 0.93 0.51 0.91 0. SDAY: Day of the year in which it is probable to happen the last frost in spring.75 -0.99 -0.40 -0.94 0.87 GSP MAP 0.79 0.37 0.91 MTWM 0.9 or higher are marked with grey.56 0.07 0.92 0.21 0.95 MMAX 0.07 0.97 0.95 0.38 -0.39 -0.09 0.85 Abutilon sp.90 0.16 -0.00 0.71 0.02 -0.50 0.89 -0.47 0.12 -0.21 -0.77 0.05 0.96 -0.69 -0.93 0.99 0. AST: Average slope of the terrain.15 0.37 -0.84 0.63 0.98 0. MTWM: Mean temperature in the warmest month.22 0.indb 295 0. Species of other genera and their covariation (C) (eq.98 0.16 0.08 -0.08 -0.95 0.95 -0.92 0.20 0.67 0.98 0.95 0.78 0.80 0.98 -0.85 0.66 -0.53 -0.62 -0.99 0.025 ordered from the highest to the lowest average absolute values (C mean).98 -0. Effect of climate and physiography on the density of tree and shrub species 295 2013-07-09 14:47:35 .99 0.84 0. GSP: Total precipitation in the growing season (April to September).24 -0.98 -0.89 -0.19 0.19 0.54 -0.78 -0.60 0.38 0.