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. 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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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

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).

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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

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.

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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:

(1)

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

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

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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.

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.

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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-

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.

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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. 5. REFERENCES
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climate change in the eastern United States Ecol. Monogr. 68: 46585. Kouchowsk i K., Teg ir mendi J. 2005 Contemporary changes of climate in Poland: Trends and variation in thermal and solar conditions related to plant vegetation Pol. J. Ecol. 59: 283297. Kronenfeld B.J., Wang Y.C h. 2007 Accounting for surveyor inconsistency and bias in estimation of tree density from presettlement land survey records Can. J. For. Res. 37: 23652379. Man ly B.F.J. 1997 Randomization, Bootstrap and Monte Carlo Methods in Biology Chapman and Hall, London, 399 pp. Maurer K., D urka W., St ck lin J. 2003 Frequency of plant species in remnants of calcareous grassland and their dispersal and persistence characteristics Basic Appl. Ecol. 4: 307316. Mirand a F., Her nnde z X.E. 1963 Los tipos de vegetacin de Mxico y su clasificacin Bol. Soc. Bot. Mx. 28: 2931. Nvar J. 2004 Preliminary testing of the ecological interactions between pines and oaks growing in mixed, unevenaged coniferous forests of Durango, Mexico Floresta, 34: 110. Pachaur i R .K., Ja l low B. 2007 Climate Change (2007): The Physical Science Basis. Working Group I, Contribution to the IPCC Fourth Assessment Report, Presentation. Nairobi, 6 February 2007. Pe ars on R .G., D aws on T.P. 2003 Predicting the impacts of climate change on the distribution of species: are bioclimate envelope models useful? Global Ecol. Biogeogr. 12: 361371. Peters on A.T., Vieg l ais D.A. 2001 Predicting Species Invasions Using Ecological Niche Modeling: New Approaches from Bioinformatics Attack a Pressing Problem BioScience, 51: 363371. R ehfeldt G.E. 2006 A spline model of climate for the western United States Gen. Tech. Rep. RMRS-GTR -165. USDA Forest Service, Fort Collins, 21 pp. R ehfeldt G.E., Cro okston N.L., War wel l M. V., Evans J.S. 2006 Empirical analyses of plant-climate relationship for the western United States Int. J. Plant. Sci. 167: 11231150.

Received after revision February 2013

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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 Pinus cembroides Zucc Pinus chihuahuana Engelm Pinus cooperi D. Don Pinus douglasiana Martnez Pinus durangensis Martnez Pinus engelmannii Carr. Pinus herrerae Martnez Freq. Density (% of (Stems/ plots) ha) Species Quercus crassifolia Quercus depressipes Trel. Quercus durifolia Seemen Quercus eduardii Trel. Quercus emoryi Quercus fulva Liebm. Quercus gentryi Muell. Quercus grisea Liebm. Quercus laeta Freq. Density (% of (Stems/ plots) ha) 17.90 15.39

0.39 0.11

0.38 0.18 0.54 1.39 0.43 0.43 0.76 1.39 0.51

14.63 11.62 Humb. & Bonpl. 9.04 7.43 2.44 3.87 7.12 1.12

0.78 10.53 8.31 3.33 5.10 2.05

0.79 5.97 5.08 2.39 2.34 0.55

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

26.00 27.52 Torr. 17.85

6.04

7.04
4.76 14.28

Pinus leiophylla Schl. 22.51 & Cham. Pinus lumholtzii Robins & Ferns Pinus luzmariae

15.91 23.16 21.34 17.66 5.71 0.61 10.14 6.11 0.39 5.18

18.24 13.28 Liebm. 4.27 0.44 3.44 1.74 0.18 1.66

16.28 Prez de la Rosa 0.64 0.47

Pinus maximinoi H.E.Moore Pinus oocarpa Schiede Pinus teocote Schlecht & Cham. Populus tremuloides Michx. Prunus serotina Ehrn Pseudotsuga menziesii Mirb Quercus acutifolia Ne

Quercus magnoliifolia Ne Quercus mcvaughii Spellenb. Quercus obtusata Humb. & Bonpl. Quercus radiata

0.24

27.72 22.98 Trel.

4.93

2.69

0.41
0.18

0.83

0.43

Quercus resinosa Liebm. Quercus rugosa Ne Quercus salicifolia Ne Quercus scytophylla Liebm.

3.60 9.04 2.22 2.88

4.21 5.00 1.49 1.66

0.89 2.88 1.22

0.18 1.55 0.39

0.28 1.23

continued overleaf

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292 APPENDIX I cont.

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

Pablo Martinez-Antnez et al.

Species

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).

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

Species

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

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journal 34.indb 293

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.
AST 0.27 0.21 0.10 0.03 -0.19 0.50 -0.02 0.25 0.24 -0.07 -0.21 0.56 0.17 -0.87 0.17 0.13 0.95 -0.04 -0.72 -0.02 0.37 -0.41 0.55 -0.26 -0.09 0.29 -0.16 0.15 -0.35 -0.47 -0.05 0.39 -0.31 0.42 -0.08 0.00 0.28 0.50 -0.24 -0.43 0.35 -0.31 -0.05 -0.54 0.70 0.38 0.03 0.17 -0.72 -0.72 -0.16 -0.08 -0.07 -0.66 -0.19 0.41 -0.20 0.28 0.05 0.19 0.06 0.95 -0.34 -0.35 0.02 -0.58 0.61 0.43 0.02 0.20 -0.57 -0.59 -0.52 -0.56 -0.85 -0.79 -0.83 -0.80 -0.78 -0.69 -0.50 -0.06 -0.55 -0.19 0.44 -0.12 -0.26 0.16 0.25 -0.13 0.04 -0.26 -0.37 -0.57 -0.21 -0.16 -0.13 0.15 -0.15 -0.10 0.27 0.22 -0.90 0.07 -0.10 -0.26 0.14 0.58 0.69 -0.54 -0.46 -0.37 -0.55 -0.58 0.37 0.35 -0.55 -0.45 -0.40 -0.49 -0.51 0.70 -0.53 0.94 0.21 0.62 0.62 0.30 0.89 0.67 0.04 -0.29 0.45 0.52 0.35 0.35 0.28 -0.76 -0.74 -0.76 -0.76 -0.69 0.70 0.72 0.67 -0.63 -0.62 -0.49 -0.71 -0.74 0.77 0.78 0.75 0.84 -0.83 0.94 0.09 0.75 0.80 0.33 0.89 0.74 -0.27 -0.53 0.48 0.47 0.37 0.60 0.55 -0.02 0.06 -0.31 0.13 0.06 0.02 -0.08 -0.18 -0.28 0.29 0.06 -0.49 -0.05 0.11 -0.10 -0.11 -0.70 -0.72 0.97 0.80 0.67 0.59 0.61 0.43 0.78 0.73 -0.91 -0.92 -0.94 -0.90 -0.83 0.44 0.45 -0.16 0.82 -0.35 -0.72 -0.76 -0.74 -0.83 0.82 -0.96 -0.23 -0.73 -0.79 -0.38 -0.92 -0.76 0.19 0.54 -0.43 -0.49 -0.34 0.88 0.67 0.76 0.78 0.57 0.36 -0.66 -0.84 0.85 0.53 0.08 0.14 0.11 0.26 0.52 0.41 -0.77 -0.77 -0.70 -0.77 -0.75 0.83 0.89 -0.88 -0.91 0.66 -0.57 -0.60 -0.83 -0.77 -0.84 0.66 -0.96 -0.06 -0.77 -0.88 -0.33 -0.92 -0.72 0.14 0.33 -0.55 -0.51 -0.46 ASP 0.04 SD -0.68 MAP 0.88 GSP 0.88 MAT -0.82 MTCM MMIN MTWM MMAX -0.76 -0.67 -0.84 -0.85 ASL -0.70 SDAY -0.78 FFP 0.84 DD5 0.48 C mean 0.68 0.63 0.61 0.61 0.59 0.58 0.57 0.52 0.51 0.48 0.48 0.46 0.44 0.44 0.43 0.39 0.38 0.37 0.28 0.24 0.22

SPECIES Pinus maximinoi

Pinus ayacahuite

Pinus douglasiana

Pinus herrerae

Cupressus lusitanica

Pinus durangensis

Pinus teocote 0.18 0.52

-0.36

Pinus cooperi

-0.55

Pinus oocarpa

Pseudotsuga menziesii

Pinus cembroides 0.39 -0.33

-0.31

Pinus leiophylla

-0.34

Abies durangensis

Juniperus flaccida

Pinus arizonica -0.64 0.04 0.14 -0.01

-0.32

Juniperus deppeana

-0.32

Juniperus monticola

Pinus luzmariae

Pinus lumholtzii

Pinus chihuahuana

-0.27

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

Pinus engelmannii

*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.

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294 Pablo Martinez-Antnez et al.

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.
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 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 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 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

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

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

*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

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journal 34.indb 295

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.
ASP 0.78 0.15 0.07 0.04 -0.10 0.39 0.20 -0.09 0.72 0.67 -0.50 0.97 -0.15 0.92 0.92 -0.64 -0.41 -0.08 -0.63 -0.99 -0.59 -0.21 -0.66 0.73 -0.20 -0.83 -0.55 0.05 -0.55 0.69 0.38 -0.47 0.39 -0.67 -0.90 -0.55 -0.04 -0.71 0.75 -0.65 0.11 -0.76 -0.98 -0.62 0.00 -0.71 0.77 0.99 0.93 0.91 0.93 0.82 0.83 0.95 -0.33 0.93 0.95 0.91 -0.22 0.40 0.76 -0.78 -0.53 -0.54 -0.77 0.78 0.35 -0.28 0.92 0.95 0.90 -0.26 0.85 0.98 -0.66 -0.75 -0.54 -0.27 0.88 0.92 -0.83 0.12 -0.72 -0.89 -0.64 0.11 -0.72 0.64 -0.59 -0.17 0.39 -0.27 0.37 -0.21 0.53 -0.02 0.25 -0.19 -0.37 0.19 -0.82 -0.84 -0.81 -0.81 -0.69 0.59 0.56 0.94 0.94 0.95 0.94 0.93 -0.85 0.98 -0.99 0.49 -0.64 0.59 -0.96 0.51 -0.17 0.66 -0.90 0.38 0.08 0.74 0.75 0.90 0.92 0.93 0.88 0.77 -0.95 0.19 -0.10 0.27 -0.07 0.18 0.33 0.07 0.24 -0.01 0.52 0.08 -0.17 -0.05 0.02 0.50 0.60 0.97 0.97 0.98 0.96 0.93 -0.89 -0.07 0.79 0.89 0.84 0.75 0.81 0.93 0.93 -0.99 -0.98 -0.95 -0.96 -0.99 0.99 -0.97 0.66 -0.76 0.72 -0.99 0.45 -0.23 0.64 -0.92 0.17 -0.08 0.22 0.69 0.70 0.98 0.98 0.98 0.97 0.95 -0.98 -0.98 0.00 0.87 0.88 0.99 0.99 0.99 0.98 0.96 -0.95 -0.96 0.97 0.98 0.98 0.95 0.96 0.99 -0.99 0.98 -0.54 0.73 -0.60 0.99 -0.40 0.21 -0.63 0.94 -0.21 0.02 0.38 0.89 0.91 0.99 0.99 0.99 0.99 0.98 -0.99 -0.99 0.99 0.54 0.98 -0.40 -0.97 0.99 -0.92 -0.92 -0.91 -0.95 -0.98 0.98 0.99 0.99 0.96 0.97 0.98 0.94 0.95 0.87 -0.99 0.99 -0.70 0.72 -0.79 0.97 -0.61 0.20 -0.71 0.89 -0.13 0.08 SD MAP GSP MAT MTCM MMIN MTWM MMAX ASL SDAY FFP DD5 C mean 0.87 0.83 0.80 0.79 0.77 0.77 0.76 0.75 0.72 0.67 0.65 0.63 0.60 0.56 0.55 0.51 0.49 0.47 0.46 0.39

SPECIES

AST

Abutilon sp.

0.85

Guazuma ulmifolia

0.40

Tilia mexicana

0.62

Colubrina heteroneura

0.65

Caesalpinia pulcherrima

0.70

Acacia macracantha

0.50

Bursera coyucensis

0.63

Bursera graveolens

0.13

Populus tremuloides

0.16

Erythrina glauca

0.83

Alnus firmifolia

-0.02

Acacia farnesiana

0.15

Alnus acuminate

0.40

Acacia berlandieri

0.56

Alnus jorullensis

0.43

Prunus serotina

0.73

Arbutus xalapensis

0.23

Bursera simaruba

0.59

Arctostaphylos pungens

-0.16

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

Acacia schaffneri

-0.28

*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.

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