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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

1 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

EFFECT OF CLIMATE AND PHYSIOGRAPHY ON THE DENSITY OF TREE AND SHRUB SPECIES IN NORTHWEST MEXICO

ABSTRACT: In order to understand the en- vironmental 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 in- dividuals of 72 species, which were found in 1804 sampling plots (covering about 123,317 km 2 ) 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 Si- erra 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 ana- lyzed. Seven of the variables showed significant cor- relation (P <0.025) with at least 74% of the studied species: ‘Julian date of last spring frost’ with an aver- age 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 5 o C’ with covariation of 0.69, ‘altitude above sea level’ with C = 0.66; ‘mean temperature in the coldest month’, ‘mean tempera- ture in the warmest month’ and ‘mean annual tem- perature’, with average values of C = 0.65 for each of these last three variables. The ‘geographic orien- tation 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 spe- cific factors influence the presence of plant species (Woodward et al. 2004), including:

land relief, soil depth, altitude above sea level and geographic location (Woodward 1987, Steffen 2008). The characteristics of climate often be- come decisive factors for the presence of plants (B eg et al. 2002, Steffen 2008, Sáenz- Romero 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 (Kożuchowski and Tegirmendžić 2005). The study of climate of the last two mil- lion years has shown that the abiotic environ- ment remains in a state of non-equilibrium, and environmental conditions follow chang- ing trends with periods of different lengths (Rehfeldt 2006, Rehfeldt et al. 2006). However, plant species have not shown a sig- nificant morphological response to climate change, but instead there is evidence of dis- persion, population fragmentation and even

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extinction (Hughes 2000, Peterson and Vieglais 2001, Goparaju 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, Pachauri and Jallow 2007, Sáenz-Romero et al. 2010, C onde et al. 2011). Several authors agree that the increase in global temperature affect biodiversity at dif- ferent scales and ways, geographically shift- ing plant and animal communities and alter- ing 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 feed- ing, also have economic and social impor- tance. For example, in forest regions forestry is a major economic activity for many peo- ple (Návar 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 (González-Elizondo et al. 2004, Go - paraju 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, Mau - rer et al. (2003) found that more frequent spe- cies are also more abundant. Density of a spe- cies 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 cli- matic and physiographic variables considered important for plants (Rehfeldt et al. 2006). The study explores relationships between tree species density to climatic variables such as accumulated precipitation in the growing sea- son, 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 help- ful to define eco-physiographic areas useful to

face a possible drastic environmental change and to strengthen other technical or scientific purposes (Tchebakova et al. 2005, Aitken et al. 2008, Zhu et al. 2011).

2. MATERIALS AND METHODS

The study was conducted in the state of Durango, which has an approximate area of 123,317 km 2 , 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 26°53’ and 22°16’ North and between 102°29’ and 107°16’ West (Fig. 1). In the Sierra Madre Occidental mountain range that crosses the State there are woodlands of Pinus, Pinus- Quercus, Quercus-Pinus, small portions of temperate mesophytic forest; and on the west- ern flanks some tropical deciduous and semi deciduous forests (González-Elizondo 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 tim- ber harvesting and, in some areas forest fires occur in periods of drought (Wehenkel et al. 2011a, González-Elizondo et al. 2012b). The 1804 primary sampling units (con- glomerates) utilized in this study were estab- lished in most of the State by the National Forest Commission (CONAFOR 2009), to perform the National Inventory of Forests and Soils (INFyS 2004−2009). The name of ‘conglomerate’ was given because each of them is integrated by a group of four circular secondary sampling units of 400 m 2 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 main- ly 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 m 2 . 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 Kronenfeld and Wang (2007), the number of stems re- corded in each plot is treated in this paper as density. The whole list of studied species is presented in Appendix I, showing the fre- quency of occurrence (Maurer et al. 2003) and the mean density of each species per hect- are in the study area. For example, Pinus her- rerae 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 vari- ables considered in this work include: the av- erage 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 Re - hfeldt (2006), were obtained from the web- site 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 Ameri- ca, Guatemala, Belize and Cuba, with records from 1976 to 1990 (Crookston et al. 2008, Sáenz-Romero et al. 2010). These data are modelled using the ANUSPLIN software, de- veloped by Hutchinson (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 pre- cipitation 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, north- west 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 cold- est month, MTWM: Mean temperature in the warmest month, MMAX: Mean maximum temperature in the warm- est 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 5°C.

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286 Pablo Martinez-Antúnez et al. Fig. 1. Location of the study area. of frost-free period (FFP,

Fig. 1. Location of the study area.

of frost-free period (FFP, days), Julian date of the last frost spring (SDAY, days) and de- gree-days above 5°C (DD5, °C). All of these variables are considered critical to the full development of plants (Tchebakova et al. 2005, Rehfeldt 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: Zenith- al 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 linear- ity or a normal distribution of the data, the relationship between number of individuals per species and plot (species density) and cli- matic and physiographic variables was mea- sured by the covariation (C) described by Gregorius et al. (2007). By definition, two ordinal variables X and Y show entire covari- ation if one variable consistently increases or consistently decreases as the other variable increases. There thus exists a strictly mono- tonic relationship (but not necessarily linear) between the two variables. The covariation C

ranges between −1 and +1, where C = 1 corre- sponds to an entirely positive covariation and C = −1 to a strictly negative covariation. If its denominator is zero, C is undefined (Grego - rius et al. 2007). Formally:

C is undefined (Grego - rius et al. 2007). Formally: (1) where: C = Covariation between

(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 sum- mation of absolute values of the indicated products. In order to test the probability that the ob- served degrees of covariation C are produced solely by random events rather than directed forces, a two-sided permutation test was per- formed based on randomly chosen reassign- ments (Manly 1997). The percentages of imi- tated 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 stud- ied species have a significant correlation (P <0.025) with at least five variables of cli- mate and physiography (Appendixes II, III and IV). The species Abutilon sp., Guazuma ulmifolia, and Tilia mexicana presented the highest correlations simultaneously with sev- eral variables and also, the largest mean val- ues 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 de- grees 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 het- eroneura and Caesalpinia pulcherrima), and two trees (Tilia mexican and Bursera coyu-

censis ) of tropical forest, and a temperate for- est 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 den-

sity 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 evi- dence 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 (Appen-

dix 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 her- rerae), 6.4% for Quercus (Q. tarahumara and Q. magnoliifolia) and 55.0% for other genres (Guazuma ulmifolia, Tilia mexicana and Col- ubrina heteroneura, among others), which means that the latter group is more demand-

ing in terms of the specific temperature re-

quirements 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 Wehen-

kel

(unpublished); however, it was decided to

report about all of them, since one of the con- tributions 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. acu- minata 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, con- sidering the absolute value of C in most of

the analyzed species. Some correlations were

negative and others positive. As happened with the variables that di- rectly indicate temperature (MAT, MTCM, MMIN, MTWM and MMAX) and those re- lated to precipitation (MAP and GSP) again, the stronger effect of ASL, SDAY, FFP and

DD5 was found in the species listed in Appen-

dix IV, where five tropical forest shrubs (Abu-

tilon sp., Colubrina heteroneura, Caesalpinia pulcherrima, Erythrina glauca and Acacia ber- landieri) showed significant C values above 0.96, while Arctostaphylos pungens (a temper-

ate forest shrub) was not found to be signifi-

(a temper- ate forest shrub) was not found to be signifi- Fig. 2. Relationship between density

Fig. 2. Relationship between density of Pinus her- rerae Martínez 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 vari- ables; these species are: Quercus tarahumara, Q. magnoliifolia, Q. subspathulata, Q. resino- sa, Q. mcvaughii, Q. acutifolia and Q. aristata. But only two of the 21 conifers (Pseu- dotsuga 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 physi- ological characteristics more resistant to low temperatures than broadleaf species, and are also more resistant to frosts. The altitude (ASL) had a significant ef- fect on the density of 65.28% of the species (Appendixes II, III and IV). This behaviour is comparable to the strong effect on the den- sity of species shown by several of the climatic variables. This is understandable, since ASL is closely related to the temperature. C ortés- Castelán and Islebe (2005) and Sharma et al. (2009) reported also significant correla- tion between altitudinal gradient, micro top- ographical and soil conditions and the rich- ness 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 Miranda and Hernández (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 (Wehenkel et al. 2011b). The results found in this study are con- sistent with those found by Iverson and Prasad (1998) who reported that the abun- dance and distribution of 80 tree species is regulated by several factors, such as data soil, topography, temperature and elevation above sea level, among others. Budke et al. (2006) also reported that topography affects fertil-

ity and soil texture, suggesting that it also af- fects 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, per- haps employing techniques as the ones used by Pearson and Dawson (2003), Hamann and Wang (2006) and S chrag et al. (2008). In general, climate variables appear to af- fect 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, in- dicating that they have a greater plasticity to adapt within their distribution rank, consid- ering the variables analyzed. These results are helpful to classify species into groups, accord- ing to the degree of correlation individually shown with each of the variables considered in this study, which may have practical appli- cations.

4. CONCLUSIONS

The results of this work may be consid- ered 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 toler- ant 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 commer- cial or restoration purposes. Given the global climatic change, if the environmental vari- ables most related to the species of interest are known, this study may also contribute to planning a program for assisted plant migra- tion, by looking for similar sites for species that currently grow in areas with specific re- quirements and for those which are in danger of extinction. Overall, this work allowed de- tecting 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, conduct- ing 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 For- est Service of the United States Department of Ag- riculture (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.

 

Freq.

Density

Freq.

Density

Freq.

Density

Species

(% of

(Stems/

Species

(% of

(Stems/

Species

(% of

(Stems/

plots)

ha)

plots)

ha)

plots)

ha)

Abies

 

durangensis

Martínez

0.39

0.38

Pinus cembroides

Zucc

14.63

11.62

Quercus crassifolia Humb. & Bonpl.

17.90

15.39

Abutilon sp.

0.11

0.18

Pinus chihuahuana

9.04

3.87

Quercus depressipes

0.78

0.79

Engelm

Trel.

Acacia

berlandieri

Benth

1.05

0.54

Pinus cooperi D. Don

7.43

7.12

Quercus durifolia

Seemen

10.53

5.97

Acacia

macracantha

2.55

1.39

Pinus douglasiana

2.44

1.12

Quercus eduardii

8.31

5.08

Humb. &

Martínez

Trel.

Bonpl 1

Acacia

farnesiana (L.)

Willd

1.22

0.43

Pinus durangensis

Martínez

26.00

27.52

Quercus emoryi

Torr.

3.33

2.39

Acacia

schaffneri S.

Watson

1.27

0.43

Pinus engelmannii

Carr.

17.85

7.04

Quercus fulva

Liebm.

5.10

2.34

Alnus

 

acuminata

Kunth

1.94

0.76

Pinus herrerae

Martínez

6.04

4.76

Quercus gentryi

Muell.

2.05

0.55

Alnus

firmifolia

Fernald

1.39

1.39

Pinus leiophylla Schl. & Cham.

22.51

14.28

Quercus grisea

Liebm.

15.91

23.16

Alnus

jorullensis

Kunth

1.39

0.51

Pinus lumholtzii Robins & Ferns

18.24

13.28

Quercus laeta

Liebm.

21.34

17.66

Arbutus

 

xalapensis

Kunth 2

40.08

16.28

Pinus luzmariae

Rosa

Pérez de la

4.27

1.74

Quercus

magnoliifolia Née

5.71

6.11

Arctostaphylos

Pinus maximinoi

 

Quercus mcvaughii

pungens

Kunth

2.55

0.64

H.E.Moore

0.44

0.18

Spellenb.

0.61

0.39

Bursera

coyucensis

Bullock

1.55

0.47

Pinus oocarpa

Schiede

3.44

1.66

Quercus obtusata Humb. & Bonpl.

10.14

5.18

Bursera

graveolens

Pinus teocote

 

Quercus radiata

Trel.

 

(Kunth)

Triana &

0.22

0.24

Schlecht &

Cham.

27.72

22.98

4.93

2.69

Planch 3

Bursera

simaruba (L.)

Sarg

1.50

0.41

Populus tremuloides

Michx.

0.83

0.43

Quercus resinosa

Liebm.

3.60

4.21

Caesalpinia

pulcherrima

0.72

0.18

Prunus serotina Ehrn

0.89

0.18

Quercus rugosa Née

9.04

5.00

(L.) Sw

Colubrina

heteroneura

Griseb. Standl

0.50

0.28

Pseudotsuga

menziesii

Mirb

2.88

1.55

Quercus salicifolia

Née

2.22

1.49

Cupressus

Quercus acutifolia

Née

 

Quercus scytophylla

Liebm.

 

lusitanica

Miller

1.83

1.23

1.22

0.39

2.88

1.66

continued overleaf

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292

Pablo Martinez-Antúnez et al.

APPENDIX I – cont.

 

Freq.

Density

Freq.

Density

Freq.

Density

Species

(% of

(Stems/

Species

(% of

(Stems/

Species

(% of

(Stems/

plots)

ha)

plots)

ha)

plots)

ha)

Erythrina

0.28

0.20

Quercus aristata Hook. & Arn. Quercus arizonica Sarg.

0.67

0.59

Quercus sideroxyla Humb. & Bonpl.

34.53

34.01

glauca

Willd. 4

 

Guazuma

1.61

1.08

4.60

3.95

Quercus splendens Née

0.89

0.38

ulmifolia

Lam.

Juniperus

Quercus candicans

Née

 

Quercus

subspathulata Trel.

 

monticola

Martínez 5

0.61

0.42

1.88

0.71

0.83

0.68

Juniperus

deppeana

Steud

35.09

14.21

Quercus castanea

Née

0.67

0.48

Quercus tarahumara

Spellenb.

0.55

0.66

Juniperus flac-

4.10

2.01

Quercus chihuahuen-

5.60

10.03

Quercus urbanii

0.55

0.43

cida Schldl

sis Trel.

Trel.

Pinus arizo-

9.87

14.90

Quercus coccolobifo-

6.82

3.91

Quercus viminea

4.21

1.45

nica

Engelm

lia Trel.

Trel.

Pinus ayaca-

21.01

8.24

Quercus conzattii

1.39

0.95

Tilia mexicana

2.88

2.27

huite

Ehrenb

 

Trel.

Schldl

* 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 González-Eli - zondo (2012a): 1 Acacia macracantha (it has not been reported in Durango, it could be A. cochliacantha), 2 Arbutus xalapensis (in Durango there are at least five tree species of Arbutus with different environmental requirements, ob- viously it was considered as only one), 3 Bursera graveolens (it has not been reported in Durango, it could be B. peni- cillata) 4 Erythrina glauca (it could be E. flabelliformis) and 5 Juniperus montícola (the one in Durango is J. blancoi).

journal 34.indb 292 2013-07-09 14:47:34
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tion, 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 5°C.

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.

*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 precipita-

C mean

DD5

FFP

SDAY

ASL

MMAX

MTWM

MMIN

MTCM

MAT

GSP

MAP

SD

ASP

AST

Pseudotsuga menziesii

Cupressus lusitanica

Juniperus monticola

Juniperus deppeana

Pinus chihuahuana

Pinus engelmannii

Pinus douglasiana

Pinus durangensis

Juniperus flaccida

Abies durangensis

Pinus cembroides

Pinus maximinoi

Pinus ayacahuite

Pinus lumholtzii

Pinus luzmariae

Pinus leiophylla

Pinus arizonica

Pinus herrerae

Pinus oocarpa

Pinus cooperi

Pinus teocote

SPECIES

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

293

0.27 0.04 -0.68 0.88 0.88 -0.82 -0.76 -0.67 -0.84 -0.85 -0.70 -0.78 0.84 0.48 0.68
0.27
0.04
-0.68
0.88
0.88
-0.82
-0.76
-0.67
-0.84
-0.85
-0.70
-0.78
0.84
0.48
0.68
0.21
0.10
0.26
0.52
0.41
-0.77
-0.77
-0.70
-0.77
-0.75
0.83
0.89
-0.88
-0.91
0.63
0.53
0.03
-0.16
0.82
0.88
0.67
0.76
0.78
0.57
0.36
-0.66
-0.84
0.85
0.66
0.61
0.08
-0.19
0.43
0.78
0.73
-0.91
-0.92
-0.94
-0.90
-0.83
0.44
0.45
-0.35
-0.57
0.61
0.14
0.50
-0.02
-0.70
-0.72
0.97
0.60
0.55
0.80
0.67
0.59
0.61
-0.72
-0.60
0.59
0.11
-0.02
0.25
0.72
0.67
-0.63
-0.62
-0.49
-0.71
-0.74
0.77
0.78
-0.76
-0.83
0.58
-0.36
0.06
0.24
0.35
0.28
-0.76
-0.74
-0.76
-0.76
-0.69
0.70
0.75
-0.74
-0.77
0.57
-0.55
-0.31
-0.07
0.37
0.35
-0.55
-0.45
-0.40
-0.49
-0.51
0.70
0.84
-0.83
-0.84
0.52
0.18
0.13
-0.21
0.58
0.69
-0.54
-0.46
-0.37
-0.55
-0.58
-0.53
-0.83
0.82
0.66
0.51
0.52
0.06
0.56
0.17
0.04
-0.26
-0.37
-0.57
-0.21
-0.16
0.48
-0.31
0.02
-0.13
-0.87
-0.85
-0.79
-0.83
-0.80
-0.78
-0.69
0.21
0.09
-0.23
-0.06
0.48
-0.34
-0.08
0.15
0.17
0.13
-0.57
-0.59
-0.52
-0.56
-0.50
0.62
0.75
-0.73
-0.77
0.46
0.39
-0.18
-0.15
0.95
0.95
-0.16
-0.08
-0.04
-0.07
-0.06
0.62
0.80
-0.79
-0.88
0.44
-0.33
-0.28
-0.10
-0.34
-0.35
-0.72
-0.72
-0.72
-0.66
-0.55
0.30
0.33
-0.38
-0.33
0.44
-0.32
0.29
0.27
-0.35
-0.47
-0.05
-0.31
-0.02
-0.19
-0.19
0.89
0.89
-0.92
-0.92
0.43
-0.32
0.06
0.22
0.02
-0.05
0.39
0.35
0.37
0.41
0.44
0.67
0.74
-0.76
-0.72
0.39
-0.64
-0.49
-0.90
-0.58
-0.54
-0.31
-0.43
-0.41
-0.20
-0.12
0.04
-0.27
0.19
0.14
0.38
0.04
-0.05
0.07
0.61
0.70
0.42
0.50
0.55
0.28
-0.26
-0.29
-0.53
0.54
0.33
0.37
0.14
0.11
-0.10
0.43
0.38
-0.08
-0.24
-0.26
0.05
0.16
0.45
0.48
-0.43
-0.55
0.28
-0.27
-0.10
-0.26
0.02
0.03
0.00
-0.09
-0.16
0.19
0.25
0.52
0.47
-0.49
-0.51
0.24
-0.01
-0.11
0.14
0.20
0.17
0.28
0.29
0.15
0.06
-0.13
0.35
0.37
-0.34
-0.46
0.22
journal 34.indb 293 2013-07-09 14:47:35
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tion, 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 5°C

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: 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 precipita-

C mean

0.27

0.17

0.37

0.36

0.46

0.46

0.49

0.39

0.20

0.10

0.40

0.30

0.15

0.15

0.32

0.52

0.42

0.34

0.44

0.38

0.53

0.43

0.31

0.41

-0.34

-0.77

-0.66

-0.36

-0.45

-0.85

-0.52

-0.12

-0.29

DD5

-0.01

-0.01

0.46

0.36

0.35

0.27

0.07

0.36

0.16

0.22

0.41

0.05

0.11

0.51

0.41

-0.64

-0.38

-0.80

-0.75

-0.35

-0.20

-0.02

-0.19

-0.08

-0.08

-0.23

0.47

0.66

0.68

0.50

0.69

0.17

0.40

0.39

0.49

0.09

0.41

0.33

0.03

FFP

SDAY

-0.47

-0.57

-0.66

-0.29

-0.59

-0.60

-0.45

-0.37

-0.02

-0.42

-0.32

-0.39

-0.13

0.24

0.64

0.78

0.80

0.30

0.35

0.35

0.19

0.03

0.01

0.01

-0.54

-0.50

-0.65

-0.24

-0.30

-0.22

-0.22

-0.05

-0.03

-0.43

-0.03

-0.03

0.38

0.80

ASL

0.65

0.65

0.14

0.24

0.27

0.08

0.31

0.45

0.13

0.03

MMAX

-0.27

-0.42

-0.58

-0.59

-0.50

-0.83

-0.17

-0.26

-0.08

-0.71

-0.51

-0.53

-0.11

0.24

0.77

0.30

0.63

0.14

0.24

0.04

0.32

0.19

0.61

0.15

MTWM

-0.84

-0.57

-0.86

-0.58

-0.49

-0.83

-0.27

-0.09

-0.08

-0.31

-0.03

0.64

0.37

0.86

0.49

0.20

0.35

0.24

0.36

0.02

0.09

0.13

0.21

0.11

MMIN

-0.44

-0.77

-0.66

-0.79

-0.14

-0.17

-0.16

-0.10

-0.40

-0.05

-0.03

-0.21

0.87

0.42

0.52

0.68

0.63

0.43

0.14

0.27

0.00

0.42

0.09

0.33

MTCM

-0.56

-0.66

-0.40

-0.75

-0.23

-0.85

-0.10

-0.10

-0.22

-0.08

-0.81

0.56

0.88

0.60

0.39

0.24

0.17

0.06

0.16

0.50

0.50

0.22

0.15

0.25

MAT

-0.36

-0.62

-0.82

-0.72

-0.28

-0.70

-0.62

-0.09

-0.41

0.44

0.87

0.37

0.36

0.72

0.04

0.17

0.07

0.20

0.09

0.08

0.03

0.33

0.23

0.01

-0.84

-0.74

-0.42

-0.40

-0.35

-0.04

-0.07

-0.06

-0.61

-0.43

0.84

0.34

0.97

0.66

0.42

0.78

0.38

0.89

GSP

0.65

0.85

0.27

0.16

0.39

0.12

MAP

-0.44

-0.86

-0.62

-0.40

-0.75

-0.34

-0.41

-0.03

-0.11

0.84

0.44

0.47

0.87

0.67

0.92

0.60

0.83

0.12

0.09

0.12

0.28

0.71

0.25

0.15

-0.24

-0.26

-0.28

-0.24

-0.04

-0.07

-0.40

-0.19

-0.23

-0.03

-0.41

-0.21

-0.01

0.34

0.36

0.59

0.30

0.26

0.21

0.25

0.05

0.01

0.11

0.01

SD

-0.04

-0.07

-0.07

-0.07

-0.18

-0.03

-0.21

0.64

0.15

ASP

0.64

0.10

0.10

0.30

0.00

0.02

0.18

0.08

0.05

0.05

0.15

0.13

0.33

0.11

0.11

-0.17

-0.10

-0.08

-0.01

-0.21

-0.01

-0.41

-0.21

AST

0.26

0.48

0.59

0.70

0.63

0.55

0.53

0.04

0.41

0.21

0.15

0.05

0.53

0.03

0.11

0.41

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

Pablo Martinez-Antúnez et al.

0.72

0.71

0.68

0.68

0.67

0.61

0.55

0.85

0.90

0.91

0.93

-0.91

0.98

0.94

0.91

0.94

0.96

0.94

-0.93

0.99

0.98

-0.92

-0.94

-0.96

-0.93

0.95

-0.98

-0.97

-0.83

-0.88

-0.92

-0.93

0.86

-0.97

-0.89

0.84

0.71

-0.56

0.42

0.66

0.03

-0.56

0.87

0.88

-0.55

0.72

0.83

0.24

-0.64

0.91

0.95

-0.60

0.62

0.89

0.57

0.53

0.92

0.92

-0.69

0.54

0.85

0.54

-0.11

0.92

0.92

-0.65

0.63

0.78

0.41

0.46

0.61

0.70

0.80

0.96

0.32

0.97

0.26

0.61

0.59

0.81

0.94

0.30

0.97

0.05

0.20

0.00

-0.16

-0.24

-0.20

-0.28

-0.61

-0.12

0.22

-0.31

0.24

0.64

0.05

-0.09

0.56

0.44

0.64

0.43

0.21

0.62

0.64

journal 34.indb 294 2013-07-09 14:47:35
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tion, 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 5°C.

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: 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 precipita-

C mean

0.47

0.46

0.49

0.39

-0.71

-0.13

DD5

0.89

0.08

-0.63

-0.21

0.02

FFP

SDAY

-0.08

0.64

0.17

0.66

ASL

0.38

0.08

MMAX

-0.64

-0.72

0.64

0.11

MTWM

-0.62

-0.71

0.77

0.00

MMIN

-0.55

-0.55

0.69

0.05

MTCM

-0.66

-0.59

-0.21

0.73

MAT

-0.55

-0.04

-0.71

0.75

-0.77

-0.54

0.78

GSP

0.35

MAP

-0.78

-0.53

0.76

0.40

-0.37

-0.19

0.25

0.19

SD

-0.17

-0.05

ASP

0.02

0.08

-0.16

-0.28

AST

0.59

0.23

Arctostaphylos pungens

Arbutus xalapensis

Bursera simaruba

Acacia schaffneri

SPECIES

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

295

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

0.99

0.99

0.96

0.97

0.98

0.94

0.95

0.87

-0.99

0.99

-0.70

0.72

-0.79

 

-0.61

0.20

0.98

0.99

0.97

0.98

0.98

0.95

0.96

0.99

-0.99

0.98

-0.54

0.73

-0.60

-0.40

0.21

-0.97

-0.99

-0.96

-0.98

-0.98

-0.95

-0.96

-0.99

0.99

-0.97

0.66

-0.76

0.72

0.45

-0.23

-0.98

-0.99

-0.95

-0.98

-0.99

-0.89

-0.95

-0.85

0.98

-0.99

0.49

-0.64

0.59

0.51

-0.17

-0.95

0.98

0.96

0.95

0.75

0.93

0.77

0.93

-0.69

-0.27

0.88

0.92

-0.83

0.12

-0.72

-0.89

-0.91

0.99

0.98

0.97

0.84

0.96

0.88

0.94

-0.81

-0.54

0.83

0.91

-0.65

0.11

-0.76

-0.98

-0.40

0.99

0.99

0.98

0.93

0.98

0.93

0.95

-0.81

-0.75

0.82

0.93

-0.47

0.38

-0.20

-0.83

-0.92

0.99

0.99

0.98

0.89

0.97

0.92

0.94

-0.84

-0.66

0.93

0.99

-0.41

-0.08

-0.63

-0.99

-0.92

0.99

0.99

0.98

0.93

0.97

0.90

0.94

-0.82

-0.15

0.92

0.92

-0.64

0.39

-0.67

-0.90

0.99

0.91

0.88

0.70

0.81

0.60

0.75

0.56

-0.59

0.98

0.85

-0.26

0.90

0.95

0.92

-0.28

0.98

0.89

0.87

0.69

0.79

0.50

0.74

0.59

-0.50

0.97

0.95

-0.33

0.93

0.95

0.91

-0.22

0.54

0.38

0.00

0.22

-0.07

0.20

-0.09

0.72

0.67

-0.17

0.39

-0.27

0.37

-0.21

0.53

-0.02

0.78

0.15

0.07

0.04

-0.10

0.39

0.19

-0.10

0.27

-0.07

0.18

0.33

0.07

0.24

-0.01

0.52

0.85

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

Abutilon sp.

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

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