ABOVE-GROUND BIOMASS ACCUMULATION AND GROWTH IN A MARGINAL macrocarpa FOREST IN CENTRAL CHILE

Sergio R. Donoso, Karen Peña-Rojas, Carolina Delgado-Flores, Alejandro Riquelme and Mariangela Paratori

SUMMARY

Nothofagus macrocarpa forests are situated in the northern lim- est had an uneven-aged coppice structure, the mean age being it of Nothofagus genus distribution in South America. Data relat- 39 years. Average diameter growth rate was 0.23cm/year, height ed to their growth conditions are insufficient. Above-ground bio- growth rate was 32.3cm/year and the mean volume growth rate mass accumulation and growth were evaluated in a secondary was 0.43m3·ha-1/year, which are lower than the values indicated N. macrocarpa forest located in the northern limit of the spe- for other species of the same genus. The accumulated above cies natural distribution. Nine plots were established, and 240 ground biomass reached 35.2ton·ha-1. The low values measured sprouts of different diameter classes were harvested. The age are explained by the marginal condition of the forest as well as and volume of each sprout were determined, and tree growth in its degradation status due to anthropogenic activity. This condi- terms of diameter and height was also analyzed. Total above- tion requires of setting measures that assure the survival and ground biomass was measured in a subsample of 62 sprouts, ac- the recovery of this forest. cording to the components: stem, branches and foliage. The for-

Introduction its distribution area. Simi- actions that should be im- (DC.) F.M. Vásquez & A. larly to other Mediterranean plemented for the protection Nothofagus genus is a key Rodr. is known as “Roble de ecosystems, N. macrocarpa and monitoring of the species element in the phytogeography Santiago” and is an endemic populations have been sub- (Hechenleitner et al., 2005; of the southern hemisphere due Chilean specie. It belongs to jected to strong anthropogenic Donoso, 2007). Consequently, to its special disjointed distri- the Nothofagus group, charac- alterations, despite the fact there is a need for appropriate bution (Van Steenis, 1972), and terized by big deciduous that they are a potential biodi- and detailed information about it is the most important com- and develops in a Mediterra- versity niche for the different the species, upon which ac- ponent of the forest formations nean climate (Ramírez, 1987; communities that are living tions focused on preservation in the austral end of South Vásquez and Rodríguez, 1999) within them (Pauchard and management may be oriented. America (Gajardo, 2001). Fur- that represents the northern- Villarroel, 2002). Additionally, The aim of this study was thermore, some of the forest most distribution of the genus given its sensitivity, the spe- to quantify the amount of species show a distribution in America (Ormazábal and cies represents an outstanding above-ground biomass and to restricted to a relatively small Benoit, 1987). element for the monitoring of evaluate volume and diameter geographic area (San Martín N. macrocarpa has been global climate changes. growth in Nothofagus macro- and Ramírez, 1987). considered as a remnant spe- The current status of ac- carpa forest located at a dry The forest ecosystems in the cies. Their populations ap- quaintance with certain con- marginal site in Mediterranean central zone of Chile, where a peared thousands of years ago cepts, such as species dynam- zone of central Chile. significant part of the species under very different climatic ics and basic forest structure belonging to Nothofagus genus conditions to the current ones, relationships, with specific Materials and Methods grow, have been subjected to representing a very complex measurable parameters like intensive exploitation pressure example of what natural veg- growth rates, forest productiv- The area considered is locat- for more than 300 years. This etation dynamics has been ity, accumulated biomass, ed on “El Roble” hill (37°58´S, has resulted in a reduction of in the area (Gajardo, 2001). area and regeneration capac- 71°01´W), in the coastal moun- the area covered by the genus Currently, it grows as dis- ity, among others, is scarce tain range 75km northeast of and a deterioration of the for- jointed populations in the or does not exist. This fact Santiago urban area, or Region est (Camus, 2006). highest mountains throughout greatly impedes the different Metropolitana, Chile. The zone

KEYWORDS / Biomass Functions / Forest Dynamics / Mediterranean Forest / Received: 08/18/2009. Modified: 12/11/2009. Accepted: 12/12/2009.

Sergio R. Donoso. Forest En- versidad de Chile, PO Box UCh, Chile. e-mail: kpena@ Mariangela Paratori. Forest En- gineer, Universidad de Chile 9206, Santiago, Chile. e-mail: uchile.cl gineer, UCh, Chile. e-mail: (UCh). Doctor in Forest Engi- [email protected] Carolina Delgado-Flores. Forest [email protected] neering, Universidad de Cór- Karen Peña-Rojas. Forest En- Engineer, UCh, Chile. e-mail: doba, Spain. Professor, UCh, gineer, UCh, Chile. Doctor [email protected] Chile. Address: Facultad de in Physiology and En- Alejandro Riquelme. Biochem- Ciencias Forestales y Conser- vironment, Universidad de ist, UCh, Chile. Professor, UCh, vación de la Naturaleza, Uni- Barcelona, Spain. Professor, Chile. e-mail: [email protected]

JAN 2010, VOL. 35 Nº 01 0378-1844/10/01/065-05 $ 3.00/0 65 Acumulación de BIOmasa aérea y crecimiento en un bosque marginal de Nothofagus macrocarpa en chile central Sergio R. Donoso, Karen Peña-Rojas, Carolina Delgado-Flores, Alejandro Riquelme y Mariangela Paratori RESUMEN Los bosques de Nothofagus macrocarpa se encuentran en el mas y hojas). El bosque presentó una estructura de monte bajo límite norte de la distribución del género Nothofagus en Suda- irregular con una edad media de 39 años. El crecimiento me- mérica. El conocimiento sobre sus condiciones de crecimiento es dio anual en diámetro fue de 0,23cm/año, en altura de 32,3cm/ insuficiente. Se evaluó el crecimiento y la acumulación de bio- año y en volumen de 0,43m3·ha-1/año, valores inferiores a los masa aérea, en un bosque secundario de N. macrocarpa situado indicados para otras especies del género. La biomasa acumu- en el límite septentrional de su distribución natural. Se estable- lada alcanza a 35,2ton·ha-1. Los bajos valores determinados se cieron nueve parcelas y se cosecharon 240 vástagos a lo lar- explican por la condición marginal del bosque y el grado de go de las diferentes clases diamétricas. Se determinó la edad y degradación generado por la actividad humana. Esta condición volumen de cada vástago, así como el crecimiento en diámetro requiere definir medidas para asegurar la sobrevivencia y recu- y altura de los árboles. En una sub-muestra de 62 vástagos, se peración de este bosque. midió la biomasa aérea total y la de sus componentes (fuste, ra-

AcumulaÇÃO de BIOmaSsa aérea E creScimento eM uM bosque marginal de Nothofagus macrocarpa NO chile central Sergio R. Donoso, Karen Peña-Rojas, Carolina Delgado-Flores, Alejandro Riquelme e Mariangela Paratori RESUMO Os bosques de Nothofagus macrocarpa se encontram no limite sa aérea total e a de seus componentes (caule, ramos e folhas). norte da distribuição do gênero Nothofagus na América do Sul. O bosque apresentou uma estrutura de capim baixo irregular O conhecimento sobre suas condições de crescimento é insufi- com uma idade media de 39 anos. O crescimento médio anual ciente. Avaliou-se o crescimento e a acumulação de biomassa em diâmetro foi de 0,23cm/ano, em altura de 32,3cm/ano e em aérea, em um bosque secundário de N. macrocarpa situado no volume de 0,43m3·ha-1/ano, valores inferiores aos indicados limite setentrional de sua distribuição natural. Estabeleceram- para outras espécies do gênero. A biomassa acumulada alcança se nove lotes e se colheram 240 hastes ao longo das diferentes 35,2ton·ha-1. Os baixos valores determinados se explicam pela classes diamétricas. Determinou-se a idade e volume de cada condição marginal do bosque e o grau de degradação gerado haste, assim como o crescimento em diâmetro e altura das ár- pela atividade humana. Esta condição requer definir medidas vores. Em uma sub-amostra de 62 hastes, foi medida a biomas- para garantir a sobrevivência e recuperação deste bosque.

is included within a Nothofagus area are part of the macrocarpa formation, covering coastal batholite 996ha (Espinosa et al., 2002). conformed by gran- There is a substantial popula- ite, granodiorite and tion of trees growing on El diorite (Espinosa et Roble hill, which represents the al., 2002). Given northern limit of the species the presence of hills natural distribution (Figure 1). with steep gradients The climate is classified as exceeding 30°, the lower temperate mesothermal soils are highly sus- stenothermal semiarid Medi- ceptible to rainfall terranean type. The thermal erosion. regime is characterized by tem- The area is pre- peratures varying from a maxi- dominantly covered mum of 26.9°C in January to by scrubs and trees a minimum of 4.1°C in July. with sclerophyllous The frost-free period lasts up to leaves as well as 206 days, and there is an aver- low xerophytic and age of 13 frosts per year. The succulent scrubs, rainfall regime is characterized thorny scrubs and by an annual mean precipita- trees, and very tall Figure 1. Location of the study area tion of 656mm, a water deficit laurel-leaved trees of 897mm, and a 7-month dry (Gajardo, 1994). vegetal species growing on El N. macrocarpa formations be- season (Santibáñez and Uribe, Vegetation in the study area Roble Hill are Lithraea causti- ing outstanding (Riedemann 1993). corresponds to the Deciduous ca, Quillaja saponaria, Peumus and Aldunate, 2000). Lubert The soils are mainly of gran- Montane Forest, the Sclerophyl- boldus, Maytenus boaria, Puya and Pliscoff (2006), indicate ite origin. The three main types lous Forest, and the Sclerophyl- sp. and several cacti species, that “Roble de Santiago” forests of igneous rocks present in the lous Andean Scrub. The main with the presence of remnant belong to the vegetative strata

66 JAN 2010, VOL. 35 Nº 01 denominated Deciduous Coastal Comparisons of age, volume TABLE I Mediterranean N. macrocarpa and basal area were carried out Age distribution by diameter classes and Ribes punctatum forest. by variance analysis (ANOVA) Age (years) The study was carried out with the F-test. Significant dif- DBH Number of in a forest dominated by N. ferences were separated with (cm) sprouts Average Minimum Maximum Standard deviation macrocarpa, with a crown standard error of the means, cover close to 70% and located to evaluate least significant dif- <3 15 23 12 53 6.4 1600-1800masl, in a south-west ferences (LSD). All tests were 3.1-6.0 77 33 14 62 6.9 towards south-east exposure. evaluated at p<0.05. Statistical 6.1-9.0 80 40 10 59 8.5 A total of nine land plots of analyses were carried out using 9.1-12.0 41 47 34 63 6.5 500m2, located in three sectors, the Statgraphic statistical pack- were measured for forest char- age (Statgraphics, 2001). 12.1-15.0 20 49 26 64 4.9 acterization, considering three >15.1 7 52 46 60 4.6 randomly replicate sample plots Results and Discussion Total 240 39 10.5 per sector. The sectors were relatively near each other, sepa- Forest structure DBH: Normal diameter at 1.3m. rated by less than 800m, and the only difference among them The woodland considered same rootstock are not homoge- 5% of the samples displayed a was their exposure. A total of for this study is a secondary neous. Considering the variety, certain degree of brown stem 80 sprouts were harvested from Nothofagus macrocarpa forest age standard deviation (n=14) rotting and only 1% showed each sector. Normal diameter that has been harvested dur- is between 17.2 and 2.6 years, damages, most probably of me- at 1.3m (DBH) and total height ing the last decades, mainly for with an average of 7.8 years. chanical origin (fall of neigh- (H) were measured, and age wood extraction, as firewood The uneven-aged structure of boring trees, snow, landslides). was determined at 0.3m in each and for coal purposes. The up- the N. macrocarpa forest has The average height of sprouts stump. Based on stand tables, a per canopy is mainly dominated not been reported for other spe- is 7.8m, ranging between 2.5 total of 62 sprouts were selected by N. macrocarpa. Accompa- cies of the Nothofagus genus. and 12.7m. Normal diam- within the diameter distribution nying scrub and herb species Mollenhauer (1975) established eter (DBH) ranges from 1.3 range (1.0-22.5cm). A tree-ring were Maytenus boaria, Schinus that N. glauca forests present to 17.9cm, with an average of analysis was undertaken, ob- montanus, Ribes punctatum, even-aged coppice structures, 8.7cm ±3.5cm (Table II). taining samples from the base Calceolaria meyeniana, Bac- especially in those areas that and every meter along the stem. charis neaei, Mutisia latifolia, have been most intensively har- Volume equation Additionally, DBH growth was Solanum ligustrinum, Tanacetum vested. Santelices and Riquelme measured. The volume of each pathenium, and Berberis actina- (2007) indicated that N. ales- The model used to estimate sprout was calculated using cantha. N. macrocarpa have a sandrii coppice presents a ho- total over bark volume (TOBV) the Smalian formula, which coppice structure, where sev- mogeneous age, since there was is a local equation, where DBH requires DBH data. eral sprouts develop from each a massive uniform regeneration is the independent variable. The above-ground biomass of stump or rootstock. No primary of stumps and resulting The model was selected due to the different components (stem, forest trees were observed in the from fire, forest harvesting, and its statistical fit and because it branches and foliage) was es- area. Vegetative reproduction other large-scale alterations. In fairly represents the biological timated. Branches correspond from rootstock shoots is a com- the case of N. alpina secondary development of the trees. The to components with a basal mon characteristic of Mediterra- forests, it has been indicated following equation and statistics diameter <5cm. The anhydrous nean ecosystems (Retana et al., that, as a consequence of an- were obtained: density of the wood and bark 1992) and some species of the thropic action and natural phe- TOBV (m3)= stem samples was determined, Nothofagus genus. The coppice nomena (fire, landslides, etc.), 0.000115439×DBH (cm)2.48824 and stem biomass was esti- structure has been reported for most of the stands presented mated for each sprout as a sum N. glauca and N. alessandrii, a second-growth forest with r2 = 0.97 of both components. among others (Mollenhauer, an even-aged coppice structure F = 1717.3 (P<0.01) For leaf and branch compo- 1975; Santelices and Riquelme, (Donoso, 1993). (76) nents of each sprout, the mate- 2007). There is no mention of Wood damage status was Standard error of the estimate= rial was stored and transported N. macrocarpa in the literature, also evaluated based on tree 0.17m3 for manual separation in the mainly due to the fact that until ring analysis. Generally, in the laboratory. After separation, all few years ago, this was consid- studied forest the sprouts were According to the forest in- collected and individualized ma- ered as a variety of N. obliqua in good condition, since only ventory plots carried out within terial was oven dried at 65°C species. until constant weight. From the The trees show a high TABLE II total number of 62 sprouts se- vegetative regeneration Dasometric characteristics of the studied forest lected for above-ground biomass from rootstocks, with up estimation, 29 were harvested to 11 sprouts per stump. Forest structure variables Standard deviation Frequency distribution during winter and 33 during The forest has an average Average DBH (cm) Trees/ha summer; therefore the foliage age of 39 ±10.5 years DBH 8.7cm 3.5cm <3 72 biomass was determined only (range 10-64; Table I). Total height 11.3m 3.5m 3.1-6.0 310 for the last ones. Equations were Age dispersion character- Density 1598 sprouts/ha 293 sprouts/ha 6.1-9.0 475 developed to estimate above- izes not only the forest Density 497 stumps/ha 75 stumps/ha 9.1-12.0 452 ground biomass, where all dif- level, but also the root- Basal area 11.3m 2·ha-1 3.5m2·ha-1 12.1-15.0 210 ferent components of the sprout stock level, where the Age 39 years 10.5 years >15.1 77 were considered. ages of sprouts from the DBH: Normal diameter at 1.3m.

JAN 2010, VOL. 35 Nº 01 67 TABLE III due to anthropic activity. These lar size was 13.4kg, with the Height and diameter growth for characteristics should result in same percentages per compo- the studied forest a high competition for light, nent as those determined by DBH classes Height growth DBH growth water, and nutrients among in- Silva (1997). The forest above- (cm) (cm/year) (cm/year) dividuals (Pacheco, 2008). ground biomass obtained from The application of selective the forest inventory reached to Annual Last 2 years Annual Last 5 years cutting of the rootstocks in the 35.2ton·ha-1, the different com- average average study area produced a higher ponents showing 29.0, 5.4 and <3 21.8 4.1 0.21 0.20 availability of water, which is 0.9ton·ha-1 for stem, branches 3.1-6.0 41.7 3.1 0.18 0.16 especially apparent in those sec- and foliage, respectively. The 6.1-9.0 33.4 3.7 0.23 0.18 tors where the largest basal area total amount of biomass is 9.1-12.0 27.9 4.2 0.25 0.17 was harvested (60%). The indi- below that measured in sev- 12.1-15.0 23.7 6.0 0.29 0.20 viduals in these areas showed eral other biomass studies. >15.1 38.0 4.9 0.37 0.20 higher DBH growth (Pacheco, For Fagus sylvatica forests Average 32.3 4.1 0.23 0.18 2008). For this reason, it is it was indicated (Huet et al., DBH: Normal diameter at 1.3m. possible to state that the appli- 2004) that above-ground bio- cation of silvicultural practices mass values range from 75 to that enable an increase in light >400ton·ha-1, depending on the the studied stand, the forest ies according to site and tree and water availability could site and the age of the stand. volume was 52.0m3·ha-1. This ages. Values range from 0.3 to lead to an increase in diameter Higher values are reported value is very low compared to 0.8cm/year, and even values up and height growth rates. for Nothofagus forests grow- the typical values observed in to 1cm/year have been found ing in Chilean Mediterranean secondary Nothofagus forests (Wadsworth, 1976; Donoso, Biomass accumulation environments. In fact, Pedrasa in Chile where, usually, volume 1988; Grosse, 1989; Donoso et (1989) estimated a total above largely exceeds 100m3·ha-1 at al., 1993; Echeverría and Lara, The models selected to esti- ground biomass of 201ton·ha-1 similar stand ages (Donoso, 2004; Santelices and Riquelme, mate total and per-component for a N. alessandri forest, and 1988; Donoso et al., 1993). 2007). However, N. glauca for- above-ground biomass were ex- Gómez (1976) estimated an ests show a similar diameter ponential equations (Table IV). amount of 351ton·ha-1 for a N. Growth growth to that of N. macro- This type of equation has been glauca forest. A review of data carpa forests. Barrales (1993) widely used by different authors carried out by Magni (1995) The sprouts show a mean an- determined growth rates rang- with an accurate fit (Schönen- indicates that biomass variation nual height growth of 32.3cm/ ing from 0.07 to 0.41cm/year, berger, 1984; Caldentey, 1995; in Nothofagus forests globally year. However, during the last with an average of 0.23cm/year. Hart et al., 2003; Cañellas et ranges from 75 to 680ton·ha-1, two years, growth rate de- Also Jara (2006) determined al., 2004). The obtained models considering natural forests with creased to 4.1cm/year (Table a growth of 0.24cm/year in a corresponded to local equations, different levels of development III). These values are below the temperate Andean secondary presenting a good correlation and human intervention. How- rates observed for other Nothof- forest. and included DBH as an inde- ever, Peri et al. (2008) estimat- agus species that grow in Med- Average volume growth pendent variable. ed an above-ground biomass of iterranean environments and during the last five years was At the sprout level, biomass 64.6ton·ha-1 in a marginal N. have a coppice structure. For 0.43m3·ha-1/year. This value is values as a function of DBH antarctica forest 40-60 years example, N. glauca, a species below the rates observed for fall within the wide range of old. The values determined that in its northernmost distri- other forests growing under values determined for different within the present study for bution shares the same area the same climatic conditions. Nothofagus formations (Calde- N. macrocarpa are below the with N. macrocarpa, shows a Higher values have been deter- ntey, 1995; Silva, 1997; Peri above mentioned reported val- mean growth rate similar to mined in forests growing under et al., 2008). For N. pumilio ues. This is explained by the those calculated within this marginal conditions. In fact, trees with a DBH of 8.9cm, fact that it grows under restric- study, of 28cm/year (Barrales, Barrales (1993) has determined the biomass determined by tive environmental conditions 1993) and 34cm/year-1 (Jara, a productivity of 4.4m3·ha-1/year Silva (1997) was 17.4kg, with that severely limit its develop- 2006). Additionally, N. obli- for N. glauca, and a similar component proportions of 83, ment. qua even-aged coppice which value was indicated by Donoso 14 and 3% for stem, branch- The studied forest is par- grows in a region with higher (1988) for N. obliqua. es and foliage, respectively. ticularly sensitive to climate precipitation conditions, displays The growth values deter- The estimated value for a N. changes (IPCC, 2007), being a growth rate of 40cm/year mined in the studied area rep- macrocarpa sprout of a simi- two the major limiting envi- (Donoso, 1988). resent the first data The mean diameter growth related to the varia- TABLE IV presented no significant dif- tion of these grow- Total and per-component biomass equations ferences (p<0.05) among the ing parameters in N. for Nothofagus macrocarpa different diameter classes (Table macrocarpa forests. III). The growth rate decreased The low growth rates Statistic parameters over the last five-year period. may reflect the geo- Biomass equations r2 E.E.E F (p<0.01) The values determined for N. graphical site condi- Foliage biomass (kg)= e (-3.3268 + 0.259742*DAP (cm)) 0.79 0.55 123.3 macrocarpa were lower than tions: low precipitation, (-2.10769 + 0.302921*DAP (cm)) those found in species of the warm summers and Branches biomass (kg)= e 0.78 0.60 219.2 (-0.0123226 + 0.27176*DAP (cm)) same genus in Chile. Annual cold winters, including Stem biomass (kg)= e 0.88 0.38 444.1 diameter growth in second- rainfall as snow and, Total biomass (kg)= e (0.154574 + 0.277754*DAP (cm)) 0.87 0.41 216.4 ary Nothofagus forests var- also, the degradation E.E.E: standard error of estimation, F: calculated value of Fisher test.

68 JAN 2010, VOL. 35 Nº 01 ronmental factors: the decrease cus pyrenaica Willd. coppice tribution of dry matter and age and biomass accumula- of the usually low level of pre- stands in Spanish central carbon biomass of Atlantic tion in marginal Nothofagus mountain. Ann. For. Sci. 61: beech (Fagus sylvatica L.) in a antarctica forest in Southern cipitations on one hand, and the 243-250. time sequience. Ann. For. Sci. Patagonia. For. Ecol. Manag. increase of summer maximal Donoso C (1993) Bosques templa- 61: 683-694. 255: 2502-2511. temperatures on the other hand dos de Chile y . Va- Hechenleitner VP, Gardner MF, Ramírez C (1987) El género (Pacheco, 2008). riación, estructura y dinámica. Thomas PI, Echeverría C, Es- Nothofagus y su importancia Editorial Universitaria. San- cobar B, Brownless P, Martínez en Chile. Bosque 8: 71-76. Conclusions tiago, Chile. 484 pp. C (2005) Plantas amenaza- Retana J, Riba J, Castell C, Espe- Donoso L (2007) Propuesta de das del centro-sur de Chile. lta JM (1992) Regeneration by intervención silvícola con fines Distribución, conservación y sprouting of holm-oak (Quer- The growth and biomass a de conservación para la for- propagación. 1 ed. Universi- cus ilex) stands exploited by values determined in the area mación boscosa de Nothofagus dad Austral de Chile y Real selection thinning. Vegetatio under study are the first data macrocarpa (Caleu, Prov. Cha- Jardín Botánico de Edimburgo. 99-100: 355-364. 188 pp. related to the behavior of these cabuco). Memoria Ingeniería Riedeman P, Aldunate A (2000) variables in N. macrocarpa Forestal. Facultad de Cien- Jara C (2006) Crecimiento de Ren- Flora Nativa de Valor Orna- cias Forestales. Universidad de ovales de Hualo Sometidos mental: Chile, Zona Centro. forests. The low growth rates Chile. Santiago, Chile. 112 pp. a Distintos Tratamientos de reflect the marginal conditions Editorial Andrés Bello. San- Donoso P (1988) Caracterización Raleo. Informe. Facultad de tiago, Chile. 134 pp. of the site: low precipitation, Ciencias Agrarias y Forestales. y proposiciones silviculturales San Martín J, Ramírez C (1987) warm summers and cold win- para renovales de roble (Nothof- Universidad Católica del Maule. Chile. 41 pp. Fitosociología de los Nothofa- ters with snow, and important agus obliqua) y raulí (Nothofa- gus de la Zona Mesomórfica degradation due to anthropic gus alpina ) en el área de pro- Luebert F, Pliscoff P (2006) Sinop- chilena. Bosque 8: 121-125 tección “Radal 7 Tazas”, VII sis Bioclimática y Vegetacional activity. Santelices R, Riquelme M (2007) Región. Bosque 9: 103-114. de Chile. Editorial Universitar- The values hereby deter- ia. Santiago, Chile. 316 pp. Antecedentes dasométricos de mined are markedly lower than Donoso P, Monfil T, Otero L, Bar- Nothofagus alessandrii de la rales L (1993) Estudio de cre- Magni C (1995) Acumulación de procedencia Coipué. Bosque those reported in other Nothofa- cimiento de plantaciones y ren- Biomasa y Contenido de Nutri- 28: 281-287. entes en Bosque Mixto Natural gus forests growing in Chilean ovales manejados de especies Santibañez F, Uribe JM (1993) De Lenga (Nothofagus pumilio Mediterranean environments, nativas en el área andina de la Atlas Agroclimático de Chile, (Poepp. et Endl.) Krasser) y provincia de Cautín y Valdivia. Regiones V y Metropolitana. and this ratifies that the studied Coihue de Magallanes (Nothof- Cien. Inv. For. 7: 253-287. Universidad de Chile - Minis- forest grows under marginal agus betuloides (Mirb.) Oerst) Echeverría C, Lara A (2004) terio de Agricultura - Fondo environmental conditions that en la XII Región. Memoria Growth patterns of secondary de Investigación Agropecu- severely limit its development. Nothofagus obliqua-Nothofa- Ingeniería Forestal. Facultad de Ciencias Agrarias y Forestales. aria - Corporación Nacional It is necessary to determine gus alpina forests in Southern de Fomento. Santiago, Chile. Chile. For. Ecol. Manag. 195: Universidad de Chile. Santiago, the optimal actions to carry in Chile. 93 pp. 65 pp. order to allow these forests to 29-43. Schönenberger W (1984) Above- Mollenhauer K (1975) Estudio Espinosa V, Moreira A, Luebert B, ground biomass of mountain survive and, more, to improve de Estructura en Renovales Espinoza G, Molina JD (2002) beech (Nothofagus solandri their capacity to adapt to global de Nothofagus glauca (Phil.) Guía de Manejo Santuario de (Hook.f.) Oerst. var.cliffortio- Krasser en Bullileo. Memoria changes. la Naturaleza Cerro el Roble. ides (Hook.f.) Poole) in differ- Ingeniería Forestal. Facultad Corporación Nacional Forestal. ent stand types near timber- de Ciencias Forestales. Uni- Santiago, Chile. 53 pp. line. NZ For. 57: 59-73. Acknowledgments versidad de Chile. Santiago, Gajardo R (1994) La Vegetación Chile. 63 pp. Silva JC (1997) Crecimiento y Natural de Chile: Clasificación Acumulación de Biomasa en This research was supported Ormazábal C, Benoit I (1987) El y Distribución Geográfica. Ed- Renovales de Coihue de Ma- by funds from the FONDECYT estado de conservación del itorial Universitaria. Santiago, gallanes (Nothofagus betu- género Nothofagus en Chile. Nº 1061100, Domeyko Bio- Chile. 165 pp. loides (Mirb.) Oerst) en el diversity Program of Univer- Bosque 8: 109-120. Gajardo R (2001) Antecedentes Sector de Río San Juan, XII Pacheco C (2008) Respuesta sity of Chile and Programa de sobre el “Roble de Santiago” Región. Memoria Ingeniería Hídrica y de Crecimiento en Bosques Mediterráneos, Fac- o “Roble blanco” (Nothofagus Forestal. 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