Forest Ecology and Management 175 (2003) 119129

Adaptive genetic variation of Prosopis chilensis (Mol) Stuntz Preliminary results from one test-site
G. Verzinoa,*, C. Carranzab,1, M. Ledesmab,1, J. Joseaua, J. Di Rienzoc,2
rdoba, C.C. 509, 5000 Cordoba, Argentina Silvicultura, Facultad de Ciencias Agropecuarias, Universidad Nacional de Co b n Forestal INTA Villa Dolores, C.C. 1, 5870 Villa Dolores, Prov. de Co rdoba, Argentina Estacio c stica y Biometr a, Facultad de Ciencias Agropecuarias, Universidad Nacional de Co rdoba, C.C. 509, 5000 Cordoba, Argentina Estad Received 26 November 2001; accepted 3 April 2002
a

Abstract Despite the potential importance of Prosopis chilensis for cultivation in the arid soils of the world, little is known about the species' pattern of adaptive genetic variation. Known patterns of adaptive genetic variation in P. chilensis could be used to implement biologically sound gene conservation and breeding programs and to provide guidelines for seed transfer for afforestation and reforestation The objectives of this work were to explore the adaptive strategy of P. chilensis (Mol) Stuntz in order to provide some orientation for the species genetic management. P. chilensis grows naturally in a large arid region of Peru, Bolivia, Chile and Argentina. In Argentina, it can be found from 258 to 348S latitude and from 648 to 698W longitude, at elevations of 4302000 m a.s.l. Seeds from 17 populations from a wide region of Argentina and from six Chilean provenances, were included in the trial. The trial was conducted in the INTA Villa Dolores Forestry Station (318540 S latitude, 658080 W longitude, 529 m a.s.l.). Seedlings were grown for 4 months in the nursery and then planted in the eld following the requirements for farm-eld tests. A completely randomized design with 23 provenances (treatments) was used. The experimental units were six-tree plots and each provenance was replicated six times. The same layout of the trial in the nursery was maintained in the eld. Plant height was measured monthly on each seedling, in the nursery, from 30 to 120 days from seeding and, in the eld, at 197, 324, 434, 498, and 505 days from seeding. The average height of the six plants in each experimental unit, for each date of measurement, was taken for statistical analysis. Principal component analysis was applied to the data matrix of average plant height, previously standardized by date of measurement, in order to differentiate proles of height growth among provenances. Multiple regression analysis was used to relate genetic variation to the environmental conditions of the seed sources. Longitude, altitude, latitude and precipitation of the seed sources are slightly associated with sensitivity to frost and initial growth rates of the plants. These results suggest the presence of weak adaptive genetic variation related to macro-environmental factors; an indication of a generalist pattern of behavior. However, they should be taken as preliminary because they were obtained from one test-site. # 2002 Elsevier Science B.V. All rights reserved.
Keywords: Prosopis chilensis; Genecology; Provenance trial; Seed transfer

Corresponding author. Tel.: 54-351-4334-105/16/17; fax: 54-351-4334-118. E-mail addresses: gverzino@agro.uncor.edu (G. Verzino), intaforestal@vdolores.com.ar (C. Carranza), intaforestal@vdolores.com.ar (M. Ledesma), jajoseau@agro.uncor.edu (J. Joseau), dirienzo@agro.uncor.edu (J. Di Rienzo). 1 Tel.: 54-354-4420-154. 2 Tel.: 54-351-4334-105/16/17. 0378-1127/02/$ see front matter # 2002 Elsevier Science B.V. All rights reserved. PII: S 0 3 7 8 - 1 1 2 7 ( 0 2 ) 0 0 1 2 4 - X

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1. Introduction The National Germplasm Bank of Prosopis of the Facultad de Ciencias Agropecuarias, Universidad rdoba (NGBP), provides seeds of Nacional de Co different Prosopis species to users all over the world. Prosopis chilensis (Mol) Stuntz, among these species, is well known in a large arid region of western Argentina, Chile, Peru, and Bolivia, because of its excellent wood quality for timber, poles, posts, rewood and charcoal, as well as for the long list of nonwoody products and services it provides (Leonardis, az et al., 1988). In Argentina, 1948; Tinto, 1977; D P. chilensis grows naturally in an area which ranges from 258 to 348S latitude and from 648 to 698W longitude, at elevations of 4302000 m a.s.l. (Fig. 1). Despite the potential importance of P. chilensis for cultivation in arid regions, little is known about the species' pattern of adaptive variation. Known patterns of adaptive variation in P. chilensis could be used to implement biologically sound gene conservation and breeding programs and to provide guidelines for seed transfer for afforestation and reforestation (Li et al., 1997b). Species adapt to environmental variations according to two contrasting strategies (Lerner, 1954; Levins, 1966; Roughgarden, 1979). One of them, referred to as ``specialization'', is the alteration of gene frequencies in response to dissimilar environments, producing genetically differentiated populations. The other strategy, referred to variously ``homeostasis'', ``phenotypic plasticity'', or ``genetic exibility'' is the capability of maintaining optimal physiological functions under a broad range of environments. When homeostasis occurs, a single genotype can produce different phenotypes depending on the environment where the species is growing. In the specialization strategy, selection discriminates against genetic variability within populations, while in the homeostatic strategy genetic variability is enhanced (Rehfeldt, 1984). P. chilensis, together with Prosopis exuosa, Prosopis alba, Prosopis nigra and Prosopis caldenia, constitutes a complex of species known as ``black and white algarrobos'' (Verga, 1995). There is no strict reproductive isolation among these ve species, so natural hybrids are rather common (Hunziker et al., 1975; Burkart, 1976a,b; Hunziker, 1986). The genetic ow among these species is of such magnitude that,

contrary to other sections in the genus, the section Algarobia displays a general genetic homogeneity with very weak genetic differentiation among populations. The absence of spatial or temporal reproductive barriers results in high within-population variability and low differentiation among populations (Verga, 1995). In morphological and isozyme studies of P. chilensis and P. exuosa (Verga, 1995), the former showed a relatively narrow genetic diversity. Verga (op.cit), based on the variability of some morphological traits, stated that the species could be considered as ``specialized''. In contrast, Julio et al. (1999) inferred that adaptation in ``algarrobos'' might be attained through phenotypic plasticity rather than by changes in gene frequencies. Genecology, the study of adaptive variation in relation to environmental gradients, can help to elucidate the extent to which P. chilensis adapts genetically to dissimilar environments. Over the last 20 years, short-term testing techniques have been developed to assay how well populations are adapted to particular environmental conditions (Rehfeldt, 1984). These 25 years tests, referred to as ``farm-eld tests'' (Carlson, 1990), are designed to provide ready detection of differentiation among populations, while traditional provenance tests require more time to achieve similar objectives (Campbell, 1986; Li et al., 1997a,b; Parker et al., 1994). By means of these tests, Pseudotsuga menziesii var glauca and Pinus contorta were described as specialists (Rehfeldt, 1979a, 1983, 1989); Pinus monticola and Larix occidentalis were classied as generalists (Rehfeldt, 1979b, 1982, 1984) while P. ponderosa was described as intermediate (Rehfeldt, 1990, 1991). The objectives of this work were to explore the adaptive strategy of P. chilensis (Mol) Stuntz in order to provide some orientation for its genetic management. 2. Materials and methods Seeds from 17 populations were collected from a wide region of Argentina in the summer of 1993 and, for the purpose of comparison, six Chilean provenances were provided by the Danida Forest Seed Centre, from a FAO/CIRF project (Fig. 1).

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Fig. 1. Location of the provenances included in the trial.

Characterization of the populations in terms of latitude, longitude, altitude, and annual rain is shown in Table 1. Maps of the species geographical distribution provided by the NGBP and the Instituto Argentino n de Zonas Aridas (IADIZA) were used de Investigacio to dene the populations to be included in the trial.

Since there were no previous guidelines as how to determine the population (provenance) limits, the following criteria were taken into account: (a) altitude: maximum of 150 m between the highest and the lowest limit of each seed collection zone (population or provenance), (b) latitude: maximum of 300 between

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Table 1 Zone, sampled trees per population, geographic, physiographic and climatic conditions of the zones Order number 1 2 3 5 6 7 8 9 10 11 12 13 15 16 17 18 19 20 21 22 23 24 25 Zone Fiambala Guandacol Talampaya Andalgala Astica Belen Villa Union Los Talas Conlara Tinogasta Chilecito Las Tucumanesas Chancani Soto Catamarca Patquia Villa Dolores Pama (Chile) Rio Pama (Chile) Pama Alto-Bajo (Chile) C.A. Monte Patria (Chile) Monte Patria (Chile) Agua Chica (Chile) Trees per population 7 13 10 11 9 10 9 10 9 9 10 10 10 8 9 9 14 8 26 10 25 7 14 Latitude (south) 278410 298300 298450 278400 308550 278320 298180 288260 318570 288030 298200 298200 258400 308510 288280 308050 318570 318090 318090 318090 308390 308420 308420 Longitude (west) 678370 688300 678590 668470 678230 678060 688140 678070 658150 678350 678300 688060 658260 648570 658470 668580 658080 718040 718040 718040 718580 708570 708570 Altitude (m a.s.l.) s 1350 1450 1120 700 1370 1350 1000 520 1300 1000 1460 460 550 546 431 529 650 900 650 450 475 475 Precipitation (mm per year) 96 96 100 308 250 411 85 83 631 163 150 71 479 500 360 250 557 161 250 162 174 150 150

the southernmost and northernmost limits of each seed collection zone. The collection, processing, and storage of genetic material were carried out according to the instructions of the First International Workshop on Genetic Resources and Germplasm Conservation of Prosopis rdoba, Argentina, 1988, unpublished). Pods were (Co collected from 7 to 10 trees in each population. Seed trees selected within a given population were separated by, at least, 100 m. Fruits of each tree were processed separately with a thresher devised specially for Prosopis pods. Samples of foliage and fruits were kept for further taxonomic verication. Equal numbers of seeds from all the trees of each population were mixed in order to make up 23 seed lots (one for each provenance). Based on the principles established by Rehfeldt (1984), the study was initially formulated to be carried out at two locations. A relatively mild environment was selected to maximize the expression of differences in growth potential and a relatively severe environment was chosen to promote the expression of variation in cold hardiness. The plantation in the

severe environment was almost completely lost due to operational constraints. Although seriously affected, the trial was maintained and precise measurements were performed on the remaining plantation to obtain, after the statistical analysis, some valuable preliminary information which is here presented. The trial was conducted in the INTA, Villa Dolores Forestry Station (318540 S latitude, 658080 W longitude, 529 m a.s.l.). The climatic conditions for the area are: mean maximum temperature of the hottest month, 32.5 8C; mean minimum temperature of the coldest month, 5 8C; frost-free period, 200 days; annual rainfall, 630 mm, mainly in spring and summer. Seedlings were grown for 4 months in the nursery in springsummer of 1994. Polyethylene bottomless containers, 420 cm3, laid out on suspended wire netting, were used to promote natural root pruning. The containers were lled with soil from Prosopis natural forest, enriched with organic matter. Regular cultural practices (weeding, irrigation, pest control) were applied to the seedlings.

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In the fall of 1994 (120 days after seeding), seedlings were planted following the requirements for farm-eld tests (Carlson, 1990; Rehfeldt, 1984). Soil was thoroughly cultivated and weeded prior to planting. Plants were watered once a month with 2 l per plant. Competing vegetation was eliminated whenever needed. Each plant was protected against rodents' attack. A completely randomized design with 23 provenances (treatments) was used. The experimental units were six-tree plots and each provenance was replicated six times. The same layout of the trial in the nursery was maintained in the eld. Plant height, number of branches, number and characteristics of the leaves, caliper diameter and length of spines were measured in this trial, but exploratory analyses showed plant height as the trait that was best associated to the geographic and climatic conditions of the provenances. Plant height was measured on each seedling, in the nursery, at 30, 60, 90, and 120 days from seeding and, in the eld, at 197, 324, 434, 498, and 505 days from seeding. 2.1. Statistical analysis The average height of the six plants in each experimental unit (hereafter average plant height), for each date of measurement, was taken for statistical analysis. Univariate techniques, such as ANOVA or linear regression analysis, proved to be of little help to nd clear associations between average plant height and the geographic, topographic or climatic features of the seed sources. Taking into account that average plant height is a repeated measure, a multivariate analysis approach was carried on. Principal component analysis was applied to the data matrix of average plant height, previously standardized by date of measurement. This technique was intended to differentiate proles of height growth among provenances in a space of lesser dimension (Johnson and Wichern, 1998). The procedure produces linear combinations of a given set of variables that maximize, under some constraints, the variance of the resulting combinations (principal components). Multiple regression analysis was used to relate genetic variation to the geographic, physiographic, and climatic conditions of the seed sources. For this analysis, the principal components were included as

dependent variables while latitude, longitude, altitude, and annual rain of the provenances were the independent variables in the regression model. Second-order polynomial terms were tentatively added in order to model curvatures. Mallows' Cp criterion was used for model selection (Hocking, 1996). 3. Results The seedlings grew very well in the nursery, with 100% survival 4 months after seeding. Two and a half months after planting in the eld, the trial was severely damaged by frost (10 8C). The event occurred in the middle of the winter of 1994, after a week of unusual hot temperature (30 8C) while the plants were actively growing. Most of the plants resprouted the following spring. Descriptive statistics for the provenances' average plant height at nine dates of measurement is summarized in Table 2. It can be noticed that some coefcients of variation are fairly high, especially in older plants, which reveals high within-populations variability, associated, in part, with the cold damage event. 3.1. Principal component analysis The rst three principal components accounted for 83% of the variability in the data set. The rst principal component accounted for 48% of the variation, the second accounted for 24% and the third accounted for 11%. Since they were interpretable, only these principal components were analyzed. Normalized principal component coefcients, calculated from the correlation matrix of average plant height measured over time, are shown in Table 3. The magnitude and sign of these coefcients reects the relative weight of each date of measurement on the principal components' scores. Dot plots representing mean standard error for the three principal components by provenances are shown in Fig. 2ac. There are not dominant variablesthis is average plant height on each date of measurementor contrasting ones, in the composition of the rst principal component (Fig. 2a). It represents the provenances' average plant height mean, over all dates of measurement. Thus, if a provenance had a high score for the rst principal

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Table 2 Mean and coefcient of variation of total height at nine different dates from seeding Provenance Statistics Days from seeding 30 1 2 3 5 6 7 8 9 10 11 12 13 15 16 17 18 19 20 21 22 23 24 25 Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV Mean CV 17.15 9.61 17.03 15.74 18.55 4.04 16.77 4.55 18.84 6.83 17.79 9.47 16.81 8.39 18.65 5.94 17.20 7.83 16.66 5.64 19.00 19.71 16.68 8.07 18.19 14.63 16.73 9.83 17.47 7.75 21.26 10.08 17.59 6.25 17.56 6.36 18.22 4.00 17.41 5.76 15.42 13.88 16.51 11.05 16.90 9.05 60 38.53 5.85 40.09 9.96 39.29 4.14 39.16 5.35 41.51 7.50 39.30 8.17 38.44 9.46 41.53 3.23 38.77 10.60 38.63 4.76 41.54 9.08 38.87 7.15 40.21 10.68 37.05 5.99 40.60 6.51 43.33 5.93 37.98 7.63 42.34 4.29 42.41 5.25 41.03 3.44 38.70 8.52 38.79 9.02 41.17 9.51 90 48.92 6.18 51.90 8.19 48.40 7.26 51.58 8.58 50.81 8.43 49.79 9.22 48.01 10.97 52.65 3.42 46.99 13.36 48.19 5.11 51.79 13.05 49.03 10.50 48.98 9.18 46.33 6.03 48.63 2.38 53.86 8.65 46.11 10.32 57.78 3.95 56.41 6.85 55.92 5.91 52.38 9.35 51.11 6.77 54.80 8.54 120 48.62 6.91 53.08 11.39 48.84 9.97 51.37 9.92 51.78 8.41 50.19 10.48 47.73 12.31 51.95 6.12 46.78 14.36 48.05 6.48 52.40 15.47 49.33 9.51 49.22 11.79 47.30 6.40 48.69 3.21 52.01 9.83 46.38 10.81 58.03 5.05 57.77 5.93 54.27 8.90 55.13 9.52 51.52 6.31 55.70 9.82 197 54.47 8.76 54.25 14.07 52.92 12.39 56.28 8.35 56.40 3.36 53.95 8.37 50.14 11.88 58.74 8.24 50.20 10.21 51.33 8.20 57.24 11.58 51.82 12.70 56.74 10.90 54.52 10.71 51.47 3.48 58.24 12.25 48.68 15.08 65.78 8.21 65.47 4.75 63.98 4.90 58.63 9.50 62.57 12.12 61.55 8.85 324 44.13 9.20 49.86 17.07 45.57 14.15 48.44 9.33 48.19 10.53 50.97 7.48 41.72 18.00 44.74 15.36 47.70 10.72 44.62 11.56 49.54 10.79 42.45 15.71 49.35 15.04 47.61 11.78 41.93 8.15 52.14 11.34 47.82 7.37 42.54 19.82 44.82 15.62 43.69 14.89 38.37 11.37 31.98 25.62 38.15 25.23 434 77.58 5.30 81.22 17.13 78.58 12.25 84.34 15.12 83.75 8.08 93.69 12.78 76.22 11.07 82.83 12.65 94.86 11.35 86.64 8.39 87.20 12.69 72.81 13.71 85.45 14.61 85.19 9.43 79.86 12.18 88.84 7.94 86.95 8.72 74.38 12.28 89.85 10.04 84.51 6.24 65.24 13.25 71.24 20.94 74.50 20.21 498 86.67 6.41 91.25 15.91 86.61 17.47 97.11 14.68 91.28 15.07 108.20 16.98 84.97 5.99 90.00 12.97 106.17 11.09 93.11 6.99 93.95 7.14 88.31 15.11 96.03 13.84 92.16 14.27 88.45 13.02 101.11 14.22 95.61 9.03 90.97 15.30 106.76 12.23 100.26 6.80 74.39 18.41 86.78 23.60 91.92 21.04 505 87.50 7.26 91.89 19.62 93.08 15.97 95.67 11.59 91.86 16.75 103.17 16.77 81.89 16.89 92.81 14.48 104.78 15.40 93.21 11.02 93.03 13.81 86.31 12.15 94.08 15.27 85.72 16.17 91.42 7.83 109.86 13.22 89.31 7.12 89.17 12.00 111.05 13.89 102.72 13.13 88.61 16.12 96.47 18.91 89.45 20.60

G. Verzino et al. / Forest Ecology and Management 175 (2003) 119129 Table 3 Normalized coefcients for the three principal components calculated from the correlation matrix of plants' height in nine dates Total height at 30 days 60 days 90 days 120 days 197 days 324 days 434 days 498 days 505 days Proportion of variance explained (%) First component 0.31 0.41 0.39 0.37 0.35 0.25 0.25 0.32 0.31 48 Second component 0.02 0.17 0.35 0.34 0.30 0.22 0.53 0.43 0.36 24 Third component 0.61 0.24 0.11 0.16 0.30 0.47 0.01 0.27 0.38 11

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component, its performance was, on the average, above the overall mean (e.g. provenances 18, 20, 21, 22). The second principal component accounts for differences in average plant height before and after the frost. Scores in this component summarize sensi-

tivity to frost. Provenances with high values for this component (e.g. 20, 23, 24, 25) had, on the average, better growth before the frost than after the event. It should be noticed that provenances 10 and 19 which showed rapid growth after the frost, are the local sources (Fig. 2b). The third component reects how trees start growing after planting or after winter rest. Provenances with high values for this component had, on the average, better growth at the beginning of the growing season than some time afterwards (Fig. 2c). On this axis, there is a clear differentiation between the Chilean provenances, on the negative side, which grew slowly at the beginning and faster afterwards, and the Argentinean ones, which grew quickly at the beginning but were outgrown by those of the Chilean origin later. Multiple regression analysis of the three principal components on the geographic, physiographic, and climatic conditions of the provenances are summarized in Table 4. The provenances' average plant height mean, over all dates of measurement (principal component 1) was positively related to longitude and annual rain and negatively related to altitude (adjusted

Table 4 Summary of estimated regression coefcients for a model where the dependent variable was the rst, the second or the third principal component and the regressors of the altitude (m a.s.l.), latitude (south), longitude (west) and annual precipitation (mm) of the 23 seed sources of P. chilensis Estimate Standard error Lower limit of confidence interval (95%) 2535.54 0.32265 24.63902 0.53190 0.00360 0.00223 19.72928 0.25021 0.04300 0.00210 0.00624 16.08843 0.01450 0.50258 0.00057 0.00458 Upper limit of confidence interval (95%) 853.57 0.39803 73.45381 0.17746 0.00055 0.01184 0.92309 0.16481 0.33463 0.00056 0.00195 31.14440 0.28806 0.28997 0.00055 0.00145 t-Statistic p-Value

Principal component 1 (adjusted R2 0:11) Constant 1694.56 425.14 Latitude 0.03769 0.18217 Longitude 49.04642 12.33882 Longitude 0.35468 0.08959 Altitude 0.00208 0.00077 Annual rain 0.00703 0.00243 Principal component 2 (adjusted R2 Constant 9.40309 Latitude 0.04270 Longitude 0.18881 Altitude 0.00133 Annual rain 0.00410 Principal component 3 (adjusted R2 Constant 23.61642 Latitude 0.13678 Longitude 0.39628 Altitude 0.00001 Annual rain 0.00301 0:42) 5.22062 0.10491 0.07372 0.00039 0.00108 0:32) 3.80593 0.07648 0.05374 0.00028 0.00079

3.98582 0.20691 3.97497 3.95889 2.69466 2.89736 1.80114 0.40704 2.56123 3.41728 3.77880 6.20516 1.78841 7.37347 0.03639 3.81001

0.0001 0.8364 0.0001 0.0001 0.0080 0.0044 0.0739 0.6846 0.0115 0.0008 0.0002 <0.0001 0.0760 <0.0001 0.9710 0.0002

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Fig. 2. Dot plots representing mean standard error for the principal components by provenance: (a) principal component 1; (b) principal component 2 and (c) principal component 3.

R2 0:11). Sensitivity to frost (principal component 2), was positively associated with longitude and negatively associated with altitude and annual rain (adjusted R2 0:42). Principal component 3, which reveals how the trees start growing after planting or after winter rest, correlated negatively with longitude and annual rain and positively with latitude (adjusted R2 0:32). 4. Discussion Principal component analysis allowed for differentiation between Chilean and Argentinean provenances

of P. chilensis in terms of sensitivity to frost and initial growth rates of plants up to 505 days old. This response probably reects differential genetic evolution of the species due to the effect of the Andes mountains, which acts as a geographical barrier between the countries. Analogous effects were described by Rehfeldt (1978, 1984) in studies about Pseudotsuga menziessii and P. contorta. Fairly high within-population variability was detected for all recorded variables (Table 2) and some among-population genetic variation was also detected. At least part of this variation appears to follow systematic patterns that can be described by functional models. Multiple regression analysis showed positive

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dependence of provenances' average plant height on the longitude and annual precipitation of the seed source: the higher the longitude and/or annual precipitation, the higher the provenances' average height. It also revealed negative dependence on altitude: the higher the altitude, the lower the provenances' average height. Earlier in this trial, Verzino et al. (1995, 1998) reported a direct relationship between the provenances' longitude and the variable total height at 197 days from seeding (the higher the longitude, the taller the plants), and an inverse relationship between the provenances' altitude and total height at 197 days from seeding (the higher the altitude, the shorter the plants). The model accounted for a signicant p 0:01 proportion of the variance R2 0:36. It was expected that correlation between genetic variability, in terms of plant height, and environmental gradients increased with time, but after 17 months of measurements the coefcient of determination remained fairly low R2 0:11, an indication that, although the environmental conditions of the provenances account signicantly for the overall plant height variability, they explain only a small part of it. Longitude of the provenances was directly related to their sensitivity to frost: the higher the longitude, the higher the sensitivity to frost. It was also inversely related to growth rate immediately after planting or after winter rest: the higher the longitude, the lower the initial growth rate. Altitude and annual rain were inversely related to sensitivity to frost: the higher the altitude and precipitation, the lower the sensitivity to frost. Annual rain was also inversely related to the growth rate after planting or after the winter. In this case, the latitude of the provenances, instead of the altitude, was positively related to the initial growth rate. Our interpretation should be taken as preliminary due to the presence of correlation between predictors, which make it hard to identify the direct contribution of each one. For example, there was an inverse relation between latitude and altitude derived from the sampling plan: in general, provenances gathered from low latitudes were mostly from high altitudes and vice versa. As a result, the high level of signicance of altitude could be masking the effect of latitude, which appeared as not signicant (Table 4). Regarding longitude, it is clear that the signicance was given

by the Chilean provenances since the relationship was not signicant when they were excluded from the analysis. Rehfeldt (1989, 1991) included altitude, longitude and latitude in predictive models for growth of P. menziesii and Pinus ponderosa var ponderosa. Latitude and altitude of the provenances were also good predictors of growth in populations of Picea engelmannii, Thuja plicata Donn ex D. Don, and L. occidentalis (Rehfeldt, 1994a,b, 1995) but no similar effect was described for longitude. It should be noticed that due to frequent interbreeding between P. chilensis and P. exuosa, some seeds used in this trial, although taken as P. chilensis, could be of hybrid origin, with high genetic variability. This leads to the concept of ``homogametic complex'' (Grant, 1976) by which P. chilensis would not be considered a biological species by itself but as part of a biological system, together with other species, and strongly subject to genetic ux. If such is the case, the adaptive strategy of the species described as P. chilensis could hardly be differentiated from the strategy of the ``complex of species'' unless careful identication of the trees collected for the trials were made by means of molecular genetic tests. In a 7-year-old provenance trial of P. exuosa, n (2000b) described high among-population Mantova variability in morphological traits associated to geographic clines. The author also reported that amongpopulation variation in phenology was correlated with n, 2000a). the latitude of the seed source (Mantova From the present study, there is only a weak evidence that variation in average plant height among populations of P. chilensis is associated with environmental gradients. Sensitivity to frost and speed of initial growth are also slightly related to environmental conditions. 5. Conclusion The poor genetic differentiation among populations of P. chilensis, along with low correlation with environmental gradients suggest a fairly generalist behavior of this species. In other words, there is a high withinpopulation variability which allows the species to adapt to different environments without major genetic alteration. This conclusion is similar to that of Julio

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G. Verzino et al. / Forest Ecology and Management 175 (2003) 119129 Johnson, R.A., Wichern, D.W., 1998. Applied Multivariate Statistical Analysis, 4th Edition. Prentice-Hall, Upper Saddle River, NJ. lisis de Julio, N., Joseau J., Saidman, B., Vilardi, J., 1999. Ana n entre variables gene ticas y ambientales en poblaciones relacio de Prosopis chilensis (Leguminosae) de Argentina y Chile. In: tica, 126 Proceedings of the XXIX Congreso Argentino de Gene pp. Leonardis, J.R., 1948. Arboles de la Argentina.Aplicaciones de su madera. Ed. Suelo Argentino. Bs As. Lerner, I.M. (Ed.), 1954. Genetic Homeostasis. Oliver & Boyd, London. Levins, R., 1966. Evolution in Changing Environments. Princeton University Press, New York. Li, P., Beaulieu, J., Bousquet, J., 1997a. Genetic structure and patterns of genetic variation among populations in eastern white spruce (Picea glauca). Can. J. For. Res. 27 (2), 189 198. Li, P., Beaulieu, J., Daoust, G., Plourde, A., 1997b. Patterns of adaptive genetic variation in eastern white pine (Pinus strobus). Can. J. For. Res. 27 (2), 199206. n, N., 2000a. Variabilidad intraespec ca de los patrones Mantova gicos de Prosopis exuosa en la provincia togeogra fenolo n Nacional del Algarrobo, Mendoza, ca del Monte. Reunio p. 30. n, N., 2000b. Diferenciacio n morfolo gica entre poblaMantova ca ciones de Prosopis exuosa D.C. en la provincia togeogra n Nacional del Algarrobo, del Monte, Argentina. Reunio Mendoza, 38 pp. Parker, W.H., Vanniejenhuis, A., Charrette, P., 1994. Adaptive variation in Picea mariana from northwestern Ontario determined by short-term common environment tests. Can. J. For. Res. 24, 8. Rehfeldt, G., 1978. Genetic differentiation of Douglas-r populations from the northern Rocky Mountains. Ecology 59 (6), 12641270. Rehfeldt, J., 1979a. Ecological adaptations in Douglas-r (Pseudotsuga menziesii var glauca) populations I. North Idaho and Northeast Washington. Heredity 43, 383397. Rehfeldt, J., 1979b. Ecotypic differentiation in Pinus monticola in northern Idaho myth or reality. Am. Nat. 114, 627736. Rehfeldt, J., 1982. Differentiation in Larix occidentalis populations from the northern Rocky Mountains. Silvae Genet. 31, 1319. Rehfeldt, J., 1983. Adaptation of Pinus contorta populations to heterogeneous environments in northern Idaho. Can. J. For. Res. 13, 405411. Rehfeldt, J., 1984. Microevolution of conifers in the northern Rocky Mountains: a view from common gardens. In: Proceedings of the Eighth North American Forest Biology Workshop, Logan, UT, pp. 132146. Rehfeldt, J., 1989. Ecological adaptations in Douglas-r, Pseudotsuga mensiessii var glauca: a synthesis. For. Ecol. Manage. 28, 203215. Rehfeldt, J., 1990. Genetic differentiation among populations of Pinus ponderosa from the upper Colorado River Basin. Bot. Gaz. 151, 125137.

et al. (1999). At present, studies on phenological behavior are carried out in order to better describe the adaptive strategy of the species. Longitude, altitude, latitude, and precipitation of the seed sources of P. chilensis are weakly associated with sensitivity to frost and initial growth rates of the plants. These results suggest the presence of some adaptive genetic variation related to macro-environmental factors, but they should be taken as preliminary because they were obtained from one test-site. Acknowledgements The authors are indebted to Dennis Joyce for helpful orientation in the project design and revision of the n Palacios for the taxonomic manuscript, to Ramo identication of the sampled trees, to Carlos Biasutti and Alicia Perez de Pereyra for their helpful coma ments and revision of the manuscript and to Secretar cnica de la Universidad Nacional de de Ciencia y Te rdoba, Instituto Nacional de Tecnolog a AgropeCo cuaria and CAFPTA for the nancial support. References
Burkart, A., 1976a. A monograph of the genus Prosopis (Leguminosae, subfam. mimosoidae). J. Arnold Arb. 57 (3), 219249. Burkart, A., 1976b. A monograph of the genus Prosopis (Leguminosae, subfam. mimosoidae). J. Arnold Arb. 57 (4), 450455. Campbell, R., 1986. Mapped genetic variation of Douglas-r to guide seed transfer in southwest Oregon. Silvae Genet. 35, 8596. Carlson, M., 1990. Early testing and selection of lodgepole pine in British Columbia. In: IUFRO Joint Working Parties Handbook. Proceedings of the IUFRO Working Parties S2.02-05,06,12 and 14 Douglas-r, Contorta Pine, Sitka Spruce and Abies Breeding and Genetic Resources. Olympia, Washington, DC. az, R., Karlin, U., Ta rtara, E., 1988. Importancia de los Prosopis D reos en las zonas a ridas y semia ridas. Documento Prosopis arbo dito. en Argentina, Ine Grant, V., 1976. Artbildung bei Panzen. Paul Parey, Berlin. Hocking, R.R., 1996. Methods and Applications of Linear Models. Regression and Analysis of Variance. Wiley, New York. Hunziker, J., 1986. Hybridization and genetic variation of Argentina species of Prosopis. For. Ecol. Manage. 16, 301315. Hunziker, J., Poggio, L., Naranjo, C., Palacios, R., Andrada, A., 1975. Cytogenetics of some species and natural hybrids in Prosopis. J. Genet. Cytol. 17, 253262.

G. Verzino et al. / Forest Ecology and Management 175 (2003) 119129 Rehfeldt, J., 1991. A model of genetic variation for applications in gene resource management. Can. J. For. Res. 21, 14911500. Rehfeldt, J., 1994a. Adaptation of Picea engelmannii populations to the heterogeneous environments of the Intermountain West. Can. J. Bot. 72, 8. Rehfeldt, J., 1994b. Genetic structure of western red cedar populations in the Interior West. Can. J. For. Res. 24, 4. Rehfeldt, J., 1995. Genetic variation, climate models and the ecological genetics of Larix occidentalis. For. Ecol. Manage. 78, 13. Roughgarden, J. (Ed.), 1979. Theory of Population Genetics and Evolutionary Ecology: An Introduction. MacMillan, New York. n de los recursos forestales argentinos. Tinto, J.C., 1977. Utilizacio cn. For. No. 41, IFONA, Bs As. Foll. Te

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ticos en Prosopis chilensis y Verga, A., 1995. Estudios gene rido Argentino. Ph.D. P. exuosa (Mimosaceae) en el Chaco A Thesis. Universitat Gottingen. Verzino, G., Carranza, C., Ledesma, M., Joseau, M., Di Rienzo, J., a de Prosopis chilensis (Mol) Stunz dentro de 1995. Genecolog n en Argentina y Chile. Resultados su rango de distribucio cnicas Forestales del Parque preliminares. Actas II. Jornadas Te o. Sgo del Estero. Chaquen Verzino, G., Ledesma, M., Carranza, C., Joseau, M., Di Rienzo, J., a de Prosopis chilensis (Mol) Stuntz dentro de 1998. Genecolog n en Arg y Chile. Avances en el su rango de distribucio conocimiento. I. Congreso Latinoamericano IUFRO El manejo o del Siglo XXI sustentable de los Recursos forestales. Desaf (en CD), Valdivia, Chile.