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Chapter 10.

Ecology of Southern Chilean

and Argentinean Nothofagus Forests
Thomas T. Veblen, Claudio Donoso,
Thomas Kitzberger and Alan J. Rebertus

TABLE OF CONTENTS

INTRODUCTION

THE PHYSICAL ENVIRONMENT AND REGIONAL VEGETATION PATTERNS

Geology and Physiography

Climate

Regional Vegetation Patterns

NOTHOFAGUS SPECIES: GENECOLOGICAL VARIATION AND HYBRIDIZATION

DISTRIBUTION, HABITATS AND FOREST TYPES

Nothofagus obliqua

Nothofagus alpina

Nothofagus dombeyi

Nothofagus nitida

Nothofagus betuloides

Nothofagus pumilio

Nothofagus antarctica

AUTECOLOGY

Reproductive Biology

Flowering, Pollination and Seed Production

Seed Dispersal, Viability and Germination Requirements

 Seedling Establishment

Vegetative Reproduction

Tree Growth

Phenology and Longevity

Height and Radial Growth

Physiological Ecology

Response to Light

Freezing Resistance

Water Relations

ASSOCIATED BIOTA

Insects

Mammals and Birds

Pathogens and Phytoparasites

FOREST DYNAMICS AND DISTURBANCE REGIMES

Patterns of Stand Development and Regeneration Modes

Coarse-scale Natural Disturbances

Disturbances of Tectonic Origin

Disturbances Associated with Glacial Processes

Blowdown

Snow Avalanches

Fire

Dieback Phenomena: Interactions of Disturbance, Climate and

Stand Dynamics

Fine-scale Treefall Gap Dynamics

NOTHOFAGUS FOREST ECOSYSTEMS

Standing Crop - Basal Areas

Standing Crop - Wood Volumes

 Standing Crop - Weight of Biomas

Leaf Area Indices

Annual Productivity

Nutrient Cycling

IMPACTS OF MODERN HUMANS

Introduced Animals

Fire and Forest Clearance

Forest Conservation and Management

Global Change and South American Nothofagus Forests

 The Ecology of Southern Chilean
and Argentinean Nothofagus Forests

Thomas T. Veblen1, Claudio Donoso2,

Thomas Kitzberger1 and Alan J. Rebertus3

1
Department of Geography, University of Colorado, Boulder,

Colorado 80309-0260, U.S.A.

2
Departamento de Silvicultura, Universidad Austral de Chile,

C.C. 567, Valdivia, Chile

3
School of Natural Resources, University of Missouri,

Columbia, Missouri 65211, U.S.A.

INTRODUCTION

In South America the genus Nothofagus extends from c. 33oS

latitude in central Chile to 56o on Tierra del Fuego (Fig. 10.1).

This chapter reviews the ecology of Nothofagus forests in

southern Chile and Argentina (south of c. 37o) and complements

the previous chapter on Nothofagus forests in the Mediterranean-

climate region of central Chile. It includes the regions known

as the Chilean and Argentinean Lake Districts (c. 38 to 43oS),

south-central Chile (c. 38 to 43oS), the Andean-Patagonian region

(south of 37oS), and Tierra del Fuego.

In southern Chile and Argentina Nothofagus occurs in two

broad classes of forests: in mixed-species temperate rainforests

on climatically favorable sites and in pure or nearly pure

Nothofagus forests on less favorable sites. The latter

relatively species-poor forests are sometimes referred to as

Patagonian or Subantartic Nothofagus forests. Both rainforests

and subantarctic Nothofagus forests are reviewed in this chapter.

THE PHYSICAL ENVIRONMENT AND REGIONAL VEGETATION PATTERNS

Geology and Physiography

The Andes of southern Chile and southwestern Argentina are a

major determinant of climate and vegetation patterns in southern

South America. From south to north they gradually rise in mean

elevation from just over 1000 m on Tierra del Fuego to well over

3000 m at c. 35oS. At 38 to 43oS along a west-to-east transect

the physiography of southern Chile and southwestern Argentina

consists of the Coastal Cordillera, the Central Depression, the

Andes, and the Patagonian Plains. The Andes, which are still

uplifting, were extensively glaciated during the Pleistocene.

Many of the glaciated surfaces have been covered by extensive

andesitic volcanic deposits of recent origin. Glaciers scoured

large lakes in the Andean foothills, and at 41 to 42oS the

glacial ice nearly extended to the foothills of the Coastal

Cordillera and covered the northwestern sector of Chiloé Island.

Northwards of c. 43oS the Andes are paralleled by the

Coastal Cordillera and the intervening Central Depression. The

Coastal Cordillera is lower in elevation than the Andes, rarely

exceeding 1000 m in elevation. It contains the oldest rocks in

2
this region, including Paleozoic and Precambrian metamorphic

rocks, and has not been glaciated (Illies 1970). The Central

Depression is a structural depression filled with Quaternary

glacial, fluvioglacial, aeolian, and alluvial deposits and

volcanic ash.

South of c. 43o the coastal mountains become islands and the

Andes rise directly from the coast. The southwest coast of South

America is a maze of fiords where on steep, glaciated surfaces

soils are typically shallow. East of the Andean summits, the

physiography of the region is subdivided into the main Andean

cordillera, the pre-cordilleran foothills, and the vast

Patagonian Plains. At 47 to 52o the Patagonian Ice Caps cover

much of the Andes and in places extend westward to the Pacific

Ocean and eastward to the large Patagonian lakes.

Climate

The southern Andes are almost continuously bathed by the

persistent mid-latitude westerlies of the Southern Hemisphere

(Miller 1976). Orographic uplift by the coastal mountains and

the Andes result in mean annual precipitations of 3000 to over

5000 mm on the windward slopes of mountains. The coastal

mountains produce a minor rainshadow effect on the Central

Depression of south-central Chile at 40oS, where mean annual

precipitation is typically less than 2000 mm (Veblen et al.

1983). Eastward of the Andes, the rainshadow effect becomes

extremely pronounced: mean annual precipitation declines from c.

3
3000 mm to less than 800 mm over a west-to-east distance of only

50 km.

Climatic variation is generally more gradual along the

latitudinal gradient. Mean annual temperature declines from

north to south, but decreasing continentality also buffers the

higher latitudes from temperature extremes. The west coast

maritime climate of South America south of c. 37oS is

characterized by a mild temperature range and high annual

precipitation (Fig. 10.2). Along the west coast as far south as

c. 42oS, Mediterranean-type precipitation seasonality is

associated with the summer presence of a subtropical high

pressure cell in the southeastern Pacific. Further south,

however, the seasonal distribution of precipitation is relatively

uniform with stormy conditions prevailing year round. In the far

south at c. 52 to 55oS, the influence of the circum-antarctic low

pressure trough becomes more evident, and cool, windy conditions

prevail for most of the year (Fig. 10.2).

Regional Vegetation Patterns

The coarse-scale vegetation patterns of southern Chile and

southwestern Argentina mirror the north-to-south and west-to-east

climatic gradients (Fig. 10.3). Bordering the evergreen

sclerophyll shrublands and low forests of Central Chile, is the

district of warm temperate deciduous Nothofagus forest. Due to

deforestation and conversion of the land to plantations of exotic

conifers, only relatively small patches of N. glauca and N.

4
obliqua remain (Chapter 9). Temperate rainforests prevail south

of c. 37o45'S. The Valdivian rainforest occurs from c. 37o45' to

43o20'S, the North Patagonian rainforest from c. 43o20' to

47o30'S, and the Magellanic rainforest south of c. 47o30'S

(Veblen et al. 1983). Although these rainforests occur mainly in

Chile, small extensions also occur on the Argentinean side of the

Andes where the physiography permits the eastern passage of humid

air masses.

As mean annual temperatures and the length of the growing

season decline along this north-to-south gradient, so do species

richness and total biomass in these rainforests. Thus, the

Valdivian rainforest constitutes the center of maximum forest

biomass and arboreal species richness (Arroyo et al. in press).

In one-hectare stands at low elevations, 10 to 15 tree species

often co-occur (Veblen et al. 1981). Although comprised mainly

of evergreen broadleaved trees and evergreen conifers, the

Valdivian rainforest district also includes the deciduous

Nothofagus obliqua and N. alpina (synonym = N. nervosa) (Table

10.1). In contrast, the Patagonian and Magellanic rainforests

are purely evergreen. Common dominants of the Patagonian rain

forests are the evergreen Nothofagus (N. dombeyi, N. nitida,

and/or N. betuloides), which typically occur in stands associated

with fewer than five or six other angiosperm or conifer tree

species. The most characteristic tree of the Magellanic

rainforests is Nothofagus betuloides which often forms monotypic

stands or co-occurs with just a few other tree species (Table

5
10.1). The deciduous N. pumilio and N. antarctica also occur

within all three rainforest districts in areas transitional to

subalpine forests (N. pumilio) or on poorly drained sites (N.

antarctica). Bamboos (Chusquea spp.) typically dominate the

rainforest understories in the north but are absent south of c.

48oS.

Parallel to the strong west-to-east precipitation gradient

is a gradient from temperate rainforests, through cool temperate

Nothofagus forests and woodlands to the Patagonian steppe of

bunchgrasses and shrubs (Fig. 10.3). Cool temperate Nothofagus

forests and woodlands occur from c. 37o30'S southwards to c. 55oS

on Tierra del Fuego and include subalpine Andean forests as well

as drier forests eastward and in the rainshadow of the Andes.

These forests and woodlands occur most extensively on the

Argentinean side of the Andes and include stands dominated by

evergreen N. dombeyi at mesic mid-elevation sites, and by

deciduous N. pumilio and N. antarctica stands at high elevation

and/or xeric sites. In the northern part of the extensive

formation of cool temperate Nothofagus forests, the evergreen

conifers Araucaria araucana (north of c. 40o20'S) and

Austrocedrus chilensis (north of c. 44oS) occur extensively along

the mesic forest to xeric woodland portion of this trans-Andean

gradient. Further south (i.e. south of c. 47oS) this strong

west-to-east precipitation gradient is also evident, but here the

conifers are absent from the xeric eastern habitats. The far

western district (west of the Magellanic rainforest), is known as

6
Magellanic moorland and is characterized by scattered N.

betuloides, bog, and ericaceous heath. This is an area of

typically waterlogged soils and nutrient-poor substrates such as

the Andean diorite of the Chilean Archipelago (Moore 1979).

Floristically, the Nothofagus forests of southern Chile and

southwestern Argentina reflect the Cretaceous and early Tertiary

connection of South America to the southwest Pacific region via

Antarctica (Chapters 2 and 11). Many of the genera in South

American Nothofagus forests are disjunctly distributed across the

southern Pacific (e.g. Araucaria, Aristotelia, Blechnum,

Caldcluvia, Discaria, Eucryphia, Gevuina, Griselinia, Hebe,

Laurelia, Lomatia, Luzuriaga, Muehlenbeckia, Nothofagus,

Podocarpus, and Pseudopanax). In addition to this strong

Gondwanaland connection, there is also an abundant neotropical

element in Nothofagus forests reflecting the long history of

relatively continuous connections with the tropical latitudes

(Arroyo et al. in press). Examples of neotropical genera include

Azara, Chusquea, Crinodendron, Dasyphyllum, Desfontainea, Drimys,

Escallonia, Myrceugenia, Raphithamnus, Schinus, and Ugni. The

connections of southern South America to the neotropics and

former Gondwanaland have contributed to the high level of generic

diversity in this region (Arroyo et al. in press).

NOTHOFAGUS SPECIES: GENECOLOGICAL VARIATION AND HYBRIDIZATION

The Nothofagus species of southern Chile and southwestern

Argentina include the deciduous species N. alpina, N. antarctica,

7
N. obliqua, and N. pumilio, and the evergreen species N.

betuloides, N. dombeyi, and N. nitida (Fig. 10.4). Their

taxonomic characteristics are described along with those of the

Central Chilean Nothofagus in Chapter 9. The broad distributions

and diversity of habitats of these species result in a high

degree of natural variability reflecting both phenotypic

plasticity and genetic differences (Donoso 1987).

The genecological variation of N. obliqua and N. alpina are

discussed in Chapter 9. There also is substantial evidence of

the genecological variation of the Nothofagus spp. of southern

Chile and Argentina. For example, seed weight of N. dombeyi

declines gradually along a north-to-south gradient from 283,688

seeds per kilo at 36o52'S to 497,310 seeds per kilo at 40o16'S

(Ordoñez 1986). The greater seed weight of the northern

populations is suggestive of ecoclinal variation in response to

increasingly drier conditions towards the north. Greater

carbohydrate reserves in the seeds of the northern populations

permit more rapid growth of seedling roots and, hence, earlier

contact with adequate supplies of soil moisture (Donoso 1987).

Similarly, the northernmost populations of Nothofagus spp. have

higher percentages of viable seed (Ordoñez 1986).

Where the distributions of the evergreen species N. dombeyi,

N. betuloides, and N. nitida overlap (40o15' to 48oS), hybrid

populations between these species are recognized by their

consistently intermediate values for traits such as number of

stamens, leaf length, and seed weight (Donoso and Atienza 1983,

8
Donoso 1984). The extent of hybridization appears greater

between N. dombeyi and N. betuloides than between either of those

species and N. nitida (A. Premoli, pers. comm.). Using starch

gel electrophoresis, the amount and distribution of genetic

variation has been compared for the widespread N. dombeyi and N.

betuloides and the geographically restricted N. nitida by Premoli

(unpublished m.s.). N. dombeyi and N. betuloides are more

genetically variable than N. nitida; they have higher total

numbers of alleles, mean number of alleles per locus, percent of

polymorphic loci, mean expected heterozygosity, and mean total

genetic diversity. The low genetic identity of N. nitida with

the other two species, the presence of four unique alleles, and

the estimated high gene flow among N. nitida populations, imply

that it has long been separated from the other two species

(Premoli unpublished m.s.). N. obliqua and N. alpina also

hybridize where their distributions overlap at mid-elevations in

Chile and Argentina (Donoso et al. 1990).

N. antarctica displays the greatest morphological variation

among South American Nothofagus (Ramírez et al. 1985, Premoli

1991). It occurs as a well formed tree 10 to 15 m tall on

optimal sites, as a small shrub (< 1 m tall) in bogs, and as a

shrubby tree (2 to 3 m tall) on rocky, xeric, and exposed sites

(e.g. timberline krummholz) (Fig. 10.5). A variety of leaf,

branch, and seed characters differ among these three morphotypes

(Ramírez et al. 1985, Premoli 1991). The genetic differences

among these strikingly different morphotypes have not yet been

9
investigated. Plants with leaf forms intermediate between N.

antarctica and N. pumilio are occasionally found but the

possibility of hybridization between these species has not yet

been formally investigated (Donoso 1987).

DISTRIBUTION, HABITATS AND FOREST TYPES

Nothofagus spp. occur in all four of the previously

described, major forest regions (i.e. Valdivian, North Patagonian

and Magellanic rainforests and the cool temperate Nothofagus

forests and woodlands). These broad forest regions can be

divided into numerous types or associations based on abundance,

cover, and basal area of dominant tree species (e.g. Yudelevich

et al. 1967, Donoso 1981, Instituto Forestal 1987) or floristic

composition (e.g. Oberdorfer 1960, Ramírez and Figueroa 1987).

However, there is no single classification widely applied to the

forests of southern Chile and southwestern Argentina. Large

areas of these forests have not been subject to any ecological

studies. In the following descriptions, for the sake of

simplicity we generally follow the forest types described for

Chile by Donoso (1981) and modify them as necessary for the

description of Nothofagus forests in Argentina.

The Nothofagus spp. of southern Chile and southwestern

Argentina generally have extensive latitudinal ranges (Fig.

10.1). Their elevational ranges all exhibit a similar pattern of

declining upper and lower limits from north to south. For

example, alpine treeline declines from c. 1800 m at 38o to 600 m

10
on Tierra del Fuego. In contrast to the more sharply delimited

plant communities of Central Chile (Chapter 9), the vegetation of

the mesic south generally follows a gradual and continuous

pattern of transition from one community type to another. Plant

species tend to be individualistically distributed, and

consequently in the following description of Nothofagus forests

only modal types are described.

Nothofagus obliqua

In Chile N. obliqua occurs from 33o to 41o30'S in the

Coastal Cordillera, in the Central Depression, and in the Andes

(Donoso 1974). In Chile south of c. 38oS, N. obliqua occurs

mostly below 600 m in the foothills of the Andes and the Coastal

Cordillera but has been largely eliminated from the Central

Depression due to forest clearance for agriculture. On the

Argentinean side of the Andes it is restricted to 36o50' to

40o15'S at elevations of 700 to 900 m (Tortorelli 1956). In the

range of N. obliqua in southern Chile and Argentina, mean annual

precipitation is 1500 to 3000 mm and the summers are usually dry.

It grows on a wide range of sites from deep, fertile soils to

relatively rocky, thin soils.

In southern Chile, the forest type in which N. obliqua is

most important is the N. obliqua-N. alpina-N. dombeyi type

(Donoso 1981). This type consists of mostly < 70-year old

second-growth stands in both the Coastal Cordillera and Andes as

far south as 40o30'S. Where it is most extensive, between 37 and

11
38oS, it extends from near sea level in the western foothills of

the coastal mountains to 1800 m in the Andes. Within this forest

type, N. obliqua increases in relative abundance towards lower

elevation and drier sites. N. dombeyi becomes more important at

moister sites and higher elevations whereas N. alpina is more

characteristic of the intermediate sites. In old-growth stands,

all three Nothofagus species occur as emergent trees attaining

heights 10 m taller than the main tree canopy at 25 to 30 m. At

low elevations, N. obliqua is more common than the other

Nothofagus spp. and codominates stands with Eucryphia cordifolia

(also an emergent tree); the other important dominant and

codominant canopy tree species include Laurelia sempervirens,

Aextoxicon punctatum, and Persea lingue. This type is

characterized by numerous species of small trees and tall shrubs.

Light gaps are typically occupied by the climbing bamboo Chusquea

quila which often has culms longer than 20 m. Due to the

suitability of the low elevation sites for agriculture, this

forest type is now rare. At c. 38oS in the Chilean Andes and

Coastal Cordillera, N. obliqua also occurs infrequently in the

lower subalpine zone (c. 1000 m), in stands dominated by N.

pumilio and Araucaria araucana.

On the Argentinean side of the Andes, N. obliqua occurs

mainly in stands codominated by N. alpina at elevations of 600 to

900 m (Fig. 10.6). Towards higher elevations and on more mesic

sites, N. dombeyi is also an important dominant. The lack of a

main canopy stratum of non-Nothofagus trees in the Argentinean

12
forests is a major contrast with the Chilean forests. At lower

elevations and towards its eastern limit, N. obliqua forms mixed

stands with the xerophytic conifer Austrocedrus chilensis.

Nothofagus alpina

N. alpina occurs in Chile between 35o and 41o30'S in both

the Andes and the Coastal Cordillera (Donoso 1993) and in

Argentina between 39o24' and 40o10'S (Tortorelli 1956).

Precipitation varies from 1500 to over 4000 mm in the range of N.

alpina in southern Chile and southwestern Argentina, and at most

sites there is a dry summer period of at least 3 months. It

typically occurs on deep and well drained soils, and is not found

in topographic depressions subject to cold air drainage.

In Chile, N. alpina occurs at low elevations in the N.

obliqua-N. alpina-N. dombeyi type previously described, which

towards higher elevations grades into the N. dombeyi-N. alpina-

Laurelia philippiana type (Donoso 1981). The latter type occurs

on moister sites from 37o to 40o30'S in the Coastal Cordillera

and Andes at 500 to 1000 m elevation. Towards its upper

elevational limit, a significant part of the precipitation falls

as snow. Soils are typically deep, fertile, and derived from

volcanic ash (Burschel et al. 1976). The Nothofagus spp. occur

as emergent trees extending at least 10 m above a dense canopy of

25 to 30-m tall Laurelia philippiana, Saxegothaea conspicua, and

Dasyphyllum diacanthoides. Light levels are low beneath the

dense tree stratum, but where gaps occur the 6 to 9-m tall bamboo

13
Chusquea culeou forms dense thickets. In Chile N. alpina also

occurs in stands that are transitional to subalpine N. pumilio

stands (Donoso 1981).

At low elevations (i.e. 600 to 800 m) on the Argentinean

side of the Andes, N. alpina is associated with N. obliqua and at

higher elevations with N. dombeyi. Most of the lower elevation

forests have been logged and sometimes converted to agriculture,

but extensive areas of N. alpina-N. dombeyi forests remain

intact. Again, these stands lack the accompanying tree species

that typify this association in Chile. The Nothofagus spp. form

relatively open canopies at heights of 35 to 45 m beneath which

the Chusquea culeou understory is well developed.

Nothofagus dombeyi

N. dombeyi occurs from 34o40' to 48oS in Chile (Donoso

1974), and from 38o30' to 44oS in Argentina (Tortorelli 1956).

It is the largest of the South American Nothofagus spp. and often

attains heights greater than 50 m and trunk diameters in excess

of 2 meters (Fig. 10.7). In elevation it ranges from near sea

level to 2500 m. In comparison with N. obliqua and N. alpina, it

occurs in moister habitats--south-facing slopes, higher

elevations, and/or riparian habitats. It is abundant on a wide

range of soil types ranging from deep, well-drained soils to

inceptisols on recent volcanic deposits and intermittently

waterlogged soils.

N. dombeyi occurs in a wide range of habitats and

14
consequently is an important component of numerous forest types.

Its associations with N. obliqua and N. alpina have already been

described. It is also an important component of the "evergreen"

or Valdivian forest type, which is highly variable but consists

exclusively of evergreen trees (Donoso 1981). This type occurs

in both the Andes and the Coastal Cordillera from 38o30' to 47oS,

from near sea level to nearly 1000 m in areas of abundant annual

precipitation (2000 to 5000 mm). The Valdivian forest type

includes a wide range of soil conditions ranging from relatively

thin podzolic soils derived from metamorphic rocks to deep well

drained volcanic soils. In the Central Depression, a variant of

the Valdivian forest type (ñadi vegetation) occurs on soils that

are intermittently waterlogged due to the development of iron-

rich hardpans in the glacially-derived sediments. Such sites are

dominated by relatively small N. dombeyi that form dense thickets

in association with the codominant Drimys winteri and abundant

myrtaceous shrubs. On well-drained soils in the Valdivian forest

type, the emergent trees are N. dombeyi, Eucryphia cordifolia,

and/or Weinmannia trichosperma, and the main canopy includes

species such as Laurelia philippiana, L. sempervirens,

Saxegothaea conspicua, Dasyphyllum diacanthoides, Drimys winteri,

Aextoxicon punctatum, Persea lingue, and/or Podocarpus nubigena

(Donoso 1981).

The Valdivian forest type extends across the Andes into

Argentina at 40 to 43oS only in a few valleys where the mean

annual precipitation is 2500 to over 3500 mm. However, there are

15
fewer tree species than on the Chilean side (typically < 4 tree

species per stand); in addition to N. dombeyi, locally codominant

species include Saxegothaea conspicua, Weinmannia trichosperma,

Podocarpus nubigena, Laurelia philippiana, Fitzroya cupressoides

and/or Dasyphyllum diacanthoides and small myrtaceous trees. At

43oS, where precipitation is highest, two to over 3-m diameter F.

cupressoides dominate this type. Bordering the Valdivian forest

type to the east, where mean annual precipitation is 1800 to 2500

mm, N. dombeyi forms extensive pure stands. These monotypic

forests have relatively open main canopies, 30 to 40 m tall,

beneath which light levels are higher than in the Chilean

rainforests (Fig. 10.8). Understories are overwhelmingly

dominated by the bamboo Chusquea culeou, which forms nearly

impenetrable 3 to 6-m tall thickets. Further east, as mean

annual precipitation declines to < 1800 mm, N. dombeyi forms

mixed stands with the xerophytic conifer Austrocedrus chilensis,

and C. culeou is sparse or absent.

Nothofagus nitida

N. nitida has a relatively restricted distribution in the

Andes and Coastal Cordillera of Chile and the southern Chilean

archipelago from c. 39o45' to 48oS (Fig. 10.9; Donoso 1974). It

occurs from near sea level to 1000 m. Climatically, its range is

characterized by mean annual precipitation > 3000 mm and cool

moist summers. N. nitida often occurs on moderately waterlogged

soils (Holdgate 1961), and generally at lower elevation than N.

16
betuloides.

N. nitida is a common component of the Valdivian and

Fitzroya cupressoides forest types (Donoso 1981) where it is

associated with N. dombeyi, N. betuloides, Fitzroya cupressoides,

Podocarpus nubigena, Saxegothaea conspicua, Eucryphia cordifolia,

Weinmannia trichosperma, Aextoxicon punctatum, Amomyrtus luma,

and Gevuina avellana. The understories of these forests are

often dominated by bamboos (Chusquea spp.) or, on poorly drained

peaty sites, by the shrub Tepualia stipularis (Holdgate 1961).

Nothofagus betuloides

N. betuloides occurs in Chile from 40o40' to 56oS (Donoso

1974, Moore 1983); in Argentina its main distribution is from 48o

to 56oS (Tortorelli 1956, Moore 1983), but we have observed

scattered occurrences close to the Chilean border at c. 43oS. It

ranges in elevation from sea level near its southern limit to as

high as 1200 m near its northern limit (Veblen et al. 1977). It

generally occurs under moister and cooler conditions and on more

waterlogged soils than N. dombeyi, and at higher elevations than

N. nitida.

North of 47oS, N. betuloides is a common component of the

Valdivian and Fitzroya cupressoides forest types (Donoso 1981).

Throughout its latitudinal range it forms an association with N.

pumilio that is transitional between wetter forests dominated by

evergreen species and drier forests dominated by N. pumilio. In

the northern part of its range (41-42oS), this association forms

17
a narrow elevational belt just below pure N. pumilio forests. In

the south (45 to 55oS), this association is transitional between

westward (coastal) pure N. betuloides forests and eastward

(interior) pure N. pumilio forests. On Tierra del Fuego N.

betuloides occurs on soils that are less fertile and poorer in

available nitrogen than the soils on which N. pumilio occurs

(Gutiérrez et al. in press). From 47o to 56oS along the coast,

on islands, and extending inland on slopes receiving high

precipitation, N. betuloides grows on the better drained soils.

These soils are typically undergoing podzolization, in contrast

to the surrounding bogs and heath vegetation where gleization

prevails (Pisano 1973, Young 1972). In coastal Magellanic

rainforests, N. betuloides typically forms a 15 to 20-m tall

canopy 5 to 8 m above an often dense stratum of Drimys winteri.

The other tall tree in this type is Podocarpus nubigena which is

locally abundant along the forest borders as far south as 51oS,

especially in areas of poor drainage. In the interior of Tierra

del Fuego, N. betuloides occurs in mid-elevation bands, bounded

above and below by N. pumilio (Rebertus and Veblen 1993b).

Nothofagus pumilio

Nothofagus pumilio occurs from 35o35' to 55oS in Chile and

Argentina in both the Andes and the Coastal Cordillera

(Tortorelli 1956, Donoso 1974). It ranges in elevation from sea

level to 2000 m. North of 41o it occurs only in the subalpine

zone and commonly forms the upper tree limit, but in the southern

18
part of its range it occurs at both high and low elevations.

Near the center of its distribution (41oS) mean annual

precipitation is > 5000 mm on the western side of the Andes and <

800 mm at the easternmost localities of N. pumilio in Argentina

(Veblen et al. 1977, Barros et al. 1983). Precipitation

generally declines towards the south so that at 55oS N. pumilio

occurs under a mean annual precipitation of 400 to 500 mm

(Tukhanen 1992). Throughout its range, much of the precipitation

falls as snow. N. pumilio occurs at cooler and drier sites than

its common assiciates N. dombeyi and N. betuloides. Thus, mixed

deciduous-evergreen Nothofagus forests give way to pure N.

pumilio forests at higher elevations and more easterly locations

in the rainshadow of the Andes (Fig. 10.10).

Although N. pumilio occurs in the Coastal Cordillera, it

usually is a component of other forest types, such as the

Araucaria type. North of c. 40oS, it is associated with N.

dombeyi, N. obliqua, N. alpina, and Araucaria araucana. South of

40o30'S in Chile and south of 48oS in Argentina, N. betuloides is

a common associate on moister sites. In the south, however, N.

pumilio typically forms extensive pure stands. North of c. 44oS,

understories are usually dominated by 1 to 3-m tall bamboo

(Chusquea tenuiflora). The bamboos are absent on dry sites, and

near timberline; south of c. 44oS the bamboos are generally

absent from the subalpine zone. At timberline, N. pumilio

krummholz often occurs in a 10 to 50-m wide ecotone with alpine

vegetation. However, at many sites, presumably those less

19
affected by strong winds, dense stands of erect N. pumilio form

timberline. Also, as described below, N. antarctica is

associated with N. pumilio.

Nothofagus antarctica

N. antarctica occurs in Chile and Argentina in the Andes and

the Coastal Cordillera from 36o30' to 56oS (Donoso 1974, Moore

1983). In elevation it ranges from near sea level in the south

to 2000 m in the north. Among South American Nothofagus, it

occurs in the widest range of habitat types: areas of poor

drainage at low to high elevations, exposed sites of unstable

substrate at alpine timberline, topographic depressions subject

to cold air drainage, steep slopes with shallow soils, and dry

sites near the Patagonian steppe ecotone (Fig. 10.5). In

general, it is common at sites which are too harsh for most other

tree species.

N. antarctica most commonly occurs in association with N.

pumilio, both at alpine timberline and at lower timberline near

the ecotone with the Patagonian steppe. At alpine timberline, N.

antarctica often occurs in krummholz vegetation either in pure

patches or interspersed with N. pumilio. Near the ecotone with

the steppe south of 45o, N. antarctica forms extensive woodlands,

sometimes intermixed with N. pumilio but generally occurring on

edaphically less favorable sites. It occurs both as a tree and

as a shrub, but the tree form is more common in the south (c. 50

to 52oS). In the north (40-43oS) on the eastern side of the

20
Andes, it forms dense thickets on steep xeric slopes with shallow

soils. In the Valdivian Rainforest Region it occurs on poorly

drained sites in ñadi vegetation in association with N. dombeyi,

and in the more hyrdric of the Fitzroya forests. It also is an

important component of the Pilgerodendron uviferum type extending

from 40o to 54oS in poorly drained areas along the Pacific coast

and at higher elevations of the Andes and Coastal Cordillera.

Other trees often found in this type are N. betuloides, Tepualia

stipularis and Drimys winteri.

AUTECOLOGY

Reproductive Biology

Flowering, Pollination and Seed Production

The southern Chile and Argentinean Nothofagus are all

monoecious and flower during spring. Flowering times vary from

September to December, with later flowering occurring at high

elevations and high latitudes (Donoso 1974). Pollen is dispersed

before female flowers are open. The nuts of Nothofagus mature

over the summer and seeds are dispersed during February to May

(Table 10.2).

Although there are no long-term studies of seed production,

field observations and several 2 to 4-year studies of seedfall

indicate that seed production and viability are highly variable

from year to year. For example, in an experimental plot in the

Andes of the Chilean Lake District, seed production of N. alpina

21
and N. dombeyi in 1970 was only 0.17 and 8.8 million seeds/ha,

respectively, but in 1971 reached 15.34 and 168.64 million

seeds/ha, respectively (Burschel et al. 1976). Percent viability

was near zero during the years of low production and ranged from

7 to 22% during the years of high production. In general for

Nothofagus, years of higher percentage viability coincide with

years of greater seed production. Seed production and viability

of N. dombeyi were consistently greater than for N. alpina

(Burschel et al. 1976).

In addition to the high annual variation in seed production,

there is significant inter-site variation. At c. 45oS in Chile,

seed production was measured for two years in two nearby pure N.

pumilio forests of similar age and at similar elevations

(Schlegel et al. 1979). Seed production over the 1977-78 period

was 0.56 and 1.44 million seeds/ha in one plot and 4.3 and 10.13

million seeds per/ha in the other. In Argentina at c. 41oS seed

rain of N. pumilio was 0.19 and 1.34 million seeds/ha in two

nearby plots in 1986 (Rusch 1993). Similarly, in Argentina at c.

40oS in two nearby stands of N. dombeyi seed rain during the 1992
season was 3.2 and 21.1 million seeds/ha (Kitzberger and Veblen

unpublished data).

Seed Dispersal, Viability and Germination Requirements

Most of the seedfall of Nothofagus occurs during a few weeks

in March to early May (Table 10.2). Germination occurs during

the spring of the same year in which the seeds are produced.

22
Germination is epigeous in which the cotyledons are elevated

above the surface by the elongating hypocotyl. Nothofagus do not

develop persistent seed banks.

Nothofagus seeds are dispersed mainly by gravity and wind.

There is much variation among species in terms of average

dispersal distance. For example, the small-seeded N. dombeyi

disperses much more effectively than the heavy-seeded N. alpina.

In a study of seed dispersal of N. pumilio in Argentina at c.

41oS Rusch (1987) found that nearly all seed fell beneath the

seed-producing tree crown. Similarly, seed dispersal

measurements at two N. dombeyi patch edges in Argentina at 40oS

indicate that 95% of the seeds fall within 20 m of the seed-

producing trees (Kitzberger and Veblen unpublished data). Air

photo comparison of the positions of these forest edges as they

advance into surrounding shrubland and grassland indicates that

95% of the expansion over a 30-year time span occurred within a

20 to 38-m radius from the initial patch edge. These results are

consistent with the norm adopted by silviculturists in southern

Chile that the distance of effective seed dispersal is

approximately one to two times the height of the tree (Donoso

1993). However, isolated seedlings of N. dombeyi found > 200 m

from the nearest possible parent tree indicate that occasionally

longer distance dispersal is achieved. N. dombeyi seeds are also

commonly found in streams which may be an effective means of

longer distance dispersal. Observations of birds infrequently

eating Nothofagus seed suggest that rare longer dispersals may be

23
possible. Furthermore, we have often inadvertently dispersed

Nothofagus seeds long distances when they are lodged in our hair

or clothing and it is likely that livestock and other animals

similarly disperse small numbers of seed.

There are numerous insects which consume Nothofagus seeds.

Perzelia spp. (Lepidoptera) consume many seeds, especially of N.

alpina. Occasionaly > 90% may be consumed and regeneration is

impeded (Carrillo and Cerda 1987). In a two year study in

Argentina, 71.4% of N. pumilio seeds were consumed by insects and

7% by rodents and litter fauna (Rusch 1987). Although birds

occasionally eat Nothofagus seeds, they account for only a tiny

percentage of the seeds consumed.

The germination behavior of South American Nothofagus spp.

is highly variable, as expected for a genus occurring in such a

broad range of habitat types. Germination requirements have been

most thoroughly investigated for N. obliqua, N. alpina, and N.

dombeyi. For these three species, the seeds of populations from

higher latitudes tend to have lower germination capacities than

lower latitude populations when exposed to the same

stratification treatment (Table 10.3). Only a few studies of the

germination behavior of other Nothofagus spp. have been

conducted. For N. betuloides at 52o11'S, 44.5% of the seed

germinated after 120 days of cold-moist stratification (Ordóñez

1986). For N. nitida at 40o12'S, only 9.5% of the seed

germinated after 120 days of cold-moist stratification (Ordóñez

1986). For N. pumilio at 52o31'S, approximately 30% of the seed

24
germinated after 20 days of cold-dry stratification (Rocuant

1984). For N. antarctica at c. 40 to 42oS, the responses to 30,

60 and 90 days of cold stratification have been shown to be

highly variable (from 0 to 18.4% germination) among populations

coming from different habitat types (Premoli 1991). In general

N. antarctica has the lowest germination capacity, but this is

offset by its vigorous vegetative reproduction (McQueen 1976).

Seedling Establishment

Given appropriate site conditions, seedling establishment of

all Nothofagus spp. can be extremely abundant. For example,

seedling (plants < 1 m tall) densities often exceed 100,000/ha

for N. pumilio and N. betuloides and > 40,000/ha for N. dombeyi

(Veblen et al. 1981, Rebertus and Veblen 1993b, Bava and Puig

1992). Seedling establishment for all Nothofagus is best under

moderately high light levels and where bare mineral soil has been

exposed. In an experimental study of seedbed preparation in a N.

dombeyi-N. alpina-Laurelia philippiana forest in the Andes at

39o35'S, Burschel et al. (1976) found that both Nothofagus

initially established abundantly where the bamboo understory and

litter cover were removed. However, nearly all seedlings died

over the subsequent 1 to 2 years due to low light levels beneath

the unaltered main canopy in combination with soil disturbance by

the ground-dwelling bird "Chucao" (Scelorchilus rubecula)

(Burschel et al. 1976).

In old-growth forests, Nothofagus seedlings often establish

25
in large numbers on fallen logs. For example, in 9 stands

sampled in the Andes of the Chilean Lake District the percentage

of seedlings established on logs ranged from 80 to 99% for N.

dombeyi, 5 to 50% for N. alpina and 10 to 87% for N. pumilio

(Veblen et al. 1981). Presence of mature N. dombeyi growing on

large decayed logs and large trees with stilt-like buttress roots

indicate that some of these seedlings survive to maturity. The

large proportion of seedlings established on logs and elevated

root buttresses may reflect poor seedling survival on the forest

floor beneath the typically dense cover of understory bamboos

(Veblen et al. 1981).

For N. pumilio, inadequate seedling establishment has been

associated with insufficient soil moisture (Rusch 1992). The

strong, desiccating winds of Patagonia can be a severe impediment

to Nothofagus regeneration in large, exposed openings (Kalela

1941, Schmidt and Urzua 1982).

The only species which develops abundant banks of persistent

'advanced growth' seedlings is N. betuloides especially at high

elevation or high latitudes where there is little competition

from understory species. On Tierra del Fuego, for example, in

old-growth N. betuloides-N. pumilio forests, N. betuloides may

persist in the understory as a seedling or sapling for 50 to 100

years (Rebertus and Veblen 1993b). In contrast, the advanced

growth of N. pumilio tends to be short-lived, but seedlings < 10-

cm tall can be abundant in undisturbed understories and can

rapidly respond to the creation of canopy gaps (Veblen et al.

26
1981, Rebertus and Veblen 1993b). Other species are either too

shade-intolerant or grow in areas with too much understory

competition to permit the development of seedling banks.

Vegetative Reproduction

Vegetative reproduction has not been reported for N. dombeyi

or N. nitida. The third evergreen species, N. betuloides has

occasionally been observed to layer in krummholz stands and to

produce basal sprouts near timberline on Tierra del Fuego

(Gutiérrez et al. 1991). All the deciduous species can reproduce

vegetatively to at least some degree. N. pumilio sometimes

produces basal shoots when damaged. For example, on Tierra del

Fuego 16.1% of 663 treefalls in 19-year old blowdowns were

reproducing by tipping (Rebertus, Veblen, and Kitzberger

unpublished). This was the dominant mode of post-blowdown

regeneration near timberline. Adventitious roots sometimes

formed where branches touched the ground, but usually new stems

relied on the roots of the original treefall.

N. antarctica depends more on vegetative reproduction than

any of the other South American Nothofagus species. It produces

root suckers (McQueen 1976) and adventitious roots on partially

buried branches (Veblen et al. 1977), and following fire it

vigorously resprouts from the base (Tortorelli 1947, Veblen and

Lorenz 1988, Burns 1993). Both N. obliqua and N. alpina

vigorously stump sprout after being cut or burned. The rapid

growth of these sprouts has resulted in extensive relatively pure

27
second-growth stands of N. obliqua and N. alpina.

Tree Growth

Phenology and Longevity.

Phenological information on the Nothofagus spp. of southern

Chile and Argentina is relatively sparse but the few phenological

studies available (Table 10.2) in combination with field

observations reveal some general patterns. In south-central

Chile, more northerly and lower elevation populations clearly

initiate leaf bud burst and flowering earlier, but there is a

high degree of variability independent of latitude and altitude

(Donoso and Cabello 1978, Becker 1981, and Mascareño 1987).

There also is a high degree of interannual variation in

phenological behavior reflecting the influence of weather (Rusch

1993). Thus, the few studies summarized in Table 2 are clearly

site and year dependent, and do not include the full range of

timing of phenological behavior of Nothofagus in Chile.

In the Argentinean Andes at c. 41o15'S, phenological

observations of N. pumilio along an elevational gradient from

1100 to 1390 m indicate a shortening of the vegetative period

with increasing elevation (Rusch 1993). At 1100 m, vegetative

growth began in September and ended in May, but was shortened at

higher elevation by delayed leaf budding and earlier leaf fall.

The diffuse-porous xylem in Nothofagus is less susceptible to

cold damage (Lechowicz 1984) and may allow species like N.

pumilio to leaf out early while frosts still occur. Leaf size

28
also declined from low to high elevations. Change in foliage

color was also simultaneous at all elevations which suggests

strong control by photoperiod. Rusch (1993) concluded that

temperature controls leaf budding, leaf size, flowering, and leaf

fall, but that photoperiod is the cue for change in foliar color.

Generally for South American Nothofagus, seedling growth

initiates earlier than the bud burst and leafing out of adult

trees. The pattern of foliar development varies substantially

among species. For example, in N. obliqua leafing out initiates

low in the canopy and progresses to the top of the canopy, but in

N. alpina the pattern is the reverse. In subalpine krummholz at

40 to 41oS in the Andes of both Argentina and Chile, N. pumilio

has been observed to initiate leaf growth a few weeks earlier

than the co-occurring N. antarctica (Eskuche 1973, Veblen et al.

1977). This may provide N. pumilio with a competitive advantage

at such sites, and, in fact, N. pumilio successionally replaces

N. antarctica in subalpine stands (Veblen et al. 1977).

N. dombeyi and N. betuloides appear to be the longest-lived

South American Nothofagus spp., occasionally attaining ages of

about 500 to 600 years (Tortorelli 1956, Lara 1991). At the

opposite extreme is N. antarctica which rarely attains ages

greater than 200 years and at ages of 80 to 120 years often

exhibits signs of senescence. Typical maximum ages for the other

species are 350 years for N. pumilio and N. nitida, and 450 years

for N. alpina and N. obliqua (LaMarche et al. 1979, Lara 1991,

Rebertus and Veblen 1993a).

29
 Height and Radial Growth.

When grown under favorable environmental conditions, the

first-year seedlings of the evergreen N. dombeyi grow

subtantially slower than N. obliqua and N. alpina. However, the

evergreen seedlings overtake the deciduous ones by the end of the

second year of development (Smulders 1988, Donoso et al. 1991a,

1991b, 1992). During the first 3 years following transplanting

of nursery-grown 2-year old seedlings, annual height increments

are higher for N. dombeyi and N. obliqua than for N. alpina

(Donoso et al. 1991a, 1991b, 1992).

Height growth of Nothofagus seedlings varies substantially

with light conditions. Under full sunlight, N. dombeyi grows

more rapidly than N. obliqua and N. alpina but the evergreen

species is the least tolerant of full shade. All three grow more

rapidly under partial shade (Müller-Using and Schlegel 1981).

Tree diameter growth also varies greatly according to site

and stand conditions. Under favorable conditions, young trees

(i.e. 25 to 70 years old) of N. dombeyi, N. obliqua and N. alpina

attain average annual diameter growth rates > 1 cm (P. Donoso

1988, P. Donoso et al. in press). In some years, diameter growth

can be > 2 cm. The diameter growth of N. pumilio is

substantially less, but in silviculturally thinned stands can

attain an annual average as high as 0.7 cm (Vera 1985).

Physiological Ecology

Although only a few ecophysiological processes have been

30
investigated (and not for all species), these initial studies

have improved the understanding of Nothofagus ecology. Given

their broad geographical ranges and high levels of morphological

and genetic variation (Premoli 1991), it is likely that different

populations of these species will have different ecophysiological

responses. Consequently, caution must be applied in generalizing

for an entire species based on ecophysiological studies of one or

a few populations.

Response to Light.

Read and Hill (1985) have compared the photosynthetic

responses of seedlings of N. alpina, N. obliqua, and N. dombeyi

(Read and Hill 1985). Photosynthetic light-dependence curves

indicate that among these species N. alpina is the most tolerant

of shade, based on its low dark respiration rate and

instantaneous light compensation point. This is consistent with

the greater abundance and height of N. alpina than N. dombeyi in

small gaps and beneath forest canopies (Veblen et al. 1981). It

is also consistent with the least shade tolerance exhibited by N.

dombeyi compared to the deciduous species in nursery studies

(Müller-Using and Schlegel 1981).

The effects of acclimation temperature (5 and 20oC) on

maximum photosynthetic rate have been investigated for seedlings

of the three evergreen Nothofagus spp. (A. Premoli, unpublished

data). N. dombeyi shifted its optimum photosynthetic temperature

much more than did N. betuloides. This may reflect the more

31
maritime distribution of the latter species, and, consequently

its exposure to a more equable temperature regime. The

geographically restricted N. nitida responded similarly to N.

dombeyi despite the former's distribution in areas of milder

temperature range. These results, in combination with patterns

of genetic diversity and the fossil record, suggest that N.

nitida formerly had a much wider distribution (Premoli in press).

Freezing Resistance.

Freezing resistances of most southern South American

Nothofagus spp. have been studied by Sakai et al. (1981), Sakai

and Wardle (1978), and Alberdi et al. (1985). N. antarctica,

followed by N. pumilio, showed the greatest tolerance for low

temperatures, which is consistent with their distributions at

high elevations, high latitudes and in frost pockets. N. obliqua

is slightly more resistant to freezing than the three evergreen

species. In general, Nothofagus spp. are less resistant to

freezing than most northern temperate trees as expected from the

more maritime climate of southern South America (Alberdi 1987).

Biochemical studies of the leaves of N. dombeyi indicate that

juvenile forms have a greater frost hardiness due to their high

content of cryoprotective carbohydrates and amino acids (Alberdi

1987). These characteristics may also enhance the ability of N.

dombeyi to colonize exposed sites.

Water Relations.

32
 Under optimal moisture availability, the transpiration rate

of N. obliqua is greater than that of N. dombeyi (Alberdi 1987).

This is consistent with the less xeromorphic leaf structure of N.

obliqua and its autumnal loss of the capacity to close stomates.

N. nitida has a lower cuticular transpiration rate than N.

dombeyi which is consistent with the greater sclerophylly (based

on fiber to protein ratios) of the former (Alberdi 1987). Lack

of daily variation in the leaf osmotic pressures of N. antarctica

and N. pumilio indicates that water content is efficiently

regulated by the opening and closing of stomates (Alberdi 1987).

Drought often results in premature leaf abscission in the

deciduous species. For example, during the dry summer of 1979 in

the Chilean Lake District N. obliqua on xeric, gravelly

substrates prematurely lost its leaves in February and March. On

the same sites, small N. dombeyi died during the drought.

Similarly, we have observed premature leaf abscission of N.

pumilio during droughts in early February 1991 on Tierra del

Fuego and in northern Patagonian Argentina (c. 41oS) in early

March 1994. In the latter area, there was a clear coincidence of

sites with early leaf abscission and lower moisture availability

as inferred from slope, aspect and soil depth.

ASSOCIATED BIOTA

Insects

There are at least 120 species of insects which feed on or

otherwise damage southern Chilean and Argentinean Nothofagus

33
(Table 10.4; Carrillo and Cerda 1987, Gentili and Gentili 1988).

Among the abundant defoliators those that occur in epidemics and

do the greatest damage are Ceropastus volupis, Hylamorpha

elegans, and Ormiscodes spp. (Carrillo and Cerda 1987). There

are numerous wood boring insects that perforate the bark of

Nothofagus and attack the wood. Such attacks by Holopterus

chilensis, Cheloderus childreni and Chilecomadia spp. facilitate

the entrance of pathogenic fungi and can render more than 50% of

the timber volume unusable. Consumers of cambium are

particularly a problem with younger trees and seedlings. There

are numerous sap suckers (especially Coccoidea) that live on

Nothofagus but they do relatively little damage. Lepidopterous

seed consumers can seriously reduce the percentage of viable

seeds.

Mammals and Birds

The mammalian fauna of South American Nothofagus forests is

usually described as sparse. This is especially true for larger

mammals, but small rodents are abundant and diverse (Pearson and

Pearson 1982). The rodents generally eat green vegetation and

fungi. Given that in Central Chile they also eat Nothofagus

seed, it is likely that they also consume tree seed in southern

Chile and Argentina.

The native herbivore with the most significant impact on

Nothofagus forests is the guanaco (Lama guanicoe), an American

camelid. Guanacos were widespread and abundant along the ecotone

34
between Nothofagus forests and the Patagonian steppe until

hunting by white settlers began at the end of the nineteenth

century. Guanacos were the principal game animal of Patagonian

aboriginals and were described by nineteenth century explorers as

occurring in immense herds (Cox 1863, Musters 1871). Although

occurring mostly in the steppe, guanacos also occurred in

adjacent Nothofagus forests (Bridges 1948, Goodall 1979) and may

have had a significant impact on these forests. Where guanacos

are abundant today, as on Tierra del Fuego, they seriously impede

Nothofagus regeneration, particularly of the deciduous species.

Other large herbivores that may have affected forest composition

include huemul deer (Hippocamelus bisulcus) and small pudu deer

(Pudu pudu). Pudu occur principally in the rainforests of

southern Chile and adjacent areas of Argentina, while the huemul

deer favors ecotonal areas near alpine timberline or the

Patagonia steppe (Povilitis 1978).

Although birds have been observed eating Nothofagus seed

(Gustavo Iglesias, pers. comm.), there are no quantitative data

on the role of Nothofagus seed in birds' diets. The majority of

tree seeds identified in the diets of birds in southern Chile and

Argentina are undigestible seeds (Armesto and Rozzi 1989).

However, large percentages of seeds and seed fragments in bird

stomachs are not identifiable but may be Nothofagus (Correa et

al. 1990). Thus, the degree to which birds utilize Nothofagus

seed is unknown.

35
Pathogens and Phytoparasites

The Nothofagus forests of southern Chile and southwestern

Argentina possess a rich mycoflora which has been the object of

much taxonomic investigation but of relatively little ecological

research (Wright 1987). The trunks of Nothofagus are attacked by

numerous pathogenic fungi such as Postia peliculosa, Piptoporus

portentosus, Ganoderma applanatum, Phlebium chrysocrea, and

Phellinus spp. (Peredo 1987, Cwielong and Rajchenberg 1992).

These and other fungi cause a high degree of trunk rot in older

trees. Gloeosoma vitellinum causes extensive decay in small

branches of N. dombeyi. Tumors on the branches of Nothofagus are

attributed to fungi such as Agrobacterium tumefaciens and Cytaria

spp. (Gamundi 1971), and the semi-parasitic plants Misodendrum

spp. (Orfila 1978). The influence of these tumors on the health

of the tree is unknown. The leaves of N. obliqua and N. alpina

are an alternate host for the fungus Mikronegeria fagi which is a

serious pest of the conifer Araucaria araucana (Butin 1969).

South American Nothofagus spp. are obligate ectotrophic

mycorrhizal, and the fungi probably enhance establishment on

bare, infertile sites (Singer and Morello 1960, Singer and Moser

1965, Singer 1971). In southern Chile and Argentinean forests

lacking Nothofagus spp. ectotrophic mycorrhizae are also lacking.

FOREST DYNAMICS AND DISTURBANCE REGIMES

Patterns of Stand Development and Regeneration Modes

36
 Patterns of stand development in South American Nothofagus

forests result from both coarse-scale disturbances producing

whole-stand replacement patterns and small treefalls resulting in

fine-scale gap dynamics. Over extensive areas in the Valdivian

rainforest district, relatively old (> 300 years old) forests are

dominated by emergent trees of shade-intolerant species such as

Nothofagus spp., Weinmannia trichosperma, and Eucryphia

cordifolia (Veblen and Ashton 1978, Veblen 1985b, Donoso et al.

1984, Donoso et al. 1985, Armesto and Figueroa 1987). In

undisturbed stands, young or small trees of these species are

absent or scarce. In contrast, shade-tolerant tree species (such

as Laurelia sempervirens, L. philippiana, Persea lingue,

Aextoxicon punctatum, and numerous myrtaceous trees) are

typically abundant and occur as all-aged populations (Veblen

1985b, Donoso et al. 1984, 1985, Armesto and Figueroa 1987). In

the absence of disturbance, there is a gradual successional trend

towards dominance by these shade-tolerant species. In landscapes

with a high frequency of coarse-scale disturbances, this

successional trend is not completed and the shade-intolerant

trees (especially Nothofagus) remain dominant in the oldest

stands (Veblen and Ashton 1978, Veblen et al. 1981). The

dependence of Nothofagus on coarse-scale disturbance is best

documented for the Valdivian rainforest district. At higher

latitudes or elevations, and on drier sites with few or no shade-

tolerant species, the tendency for Nothofagus to be

successionally replaced is obviously less or absent (Veblen

37
1989b).

In landscapes dominated by a high rate of coarse-scale

disturbance, patterns of stand development are well described by

Oliver's (1981) general model of whole-stand replacement. After

a major disturbance, Nothofagus typically pass through several

recognizable stages: (1) stem intitiation (2) stem exclusion, (3)

stem reinitiation, and (4) old growth. The model applies both to

pure or nearly pure Nothofagus forests where coarse-scale

disturbance results in re-establishment of the former canopy

dominants and to mixed-species rainforests where successional

species replacements occur. In both types of forests, the stem-

initiation and stem-exclusion stages consist of pure or nearly

pure stands of Nothofagus. Thinning is compensated by lateral

crown expansion of adjacent trees rather than new establishment

in the small gaps. In areas lacking shade-tolerant trees, N.

dombeyi, N. pumilio, and N. betuloides stands enter the stem

reinitiation phase at c. 150-200 years, when canopy gaps become

large enough to foster seedling development. If the stand is not

affected by another coarse-scale disturbance, it continues to

develop into the old-growth stage in which fine-scale treefalls

are common. In the absence of shade-tolerant tree species,

individual treefalls result in the regeneration of Nothofagus

spp.

In rainforests of Nothofagus mixed with shade-tolerant

trees, the pattern of stand development following coarse-scale

disturbance varies according to the type of disturbance and its

38
influence on understory vegetation and substrate. Disturbances

which remove nearly all the pre-existing vegetation and expose

bare mineral soil (such as mass movements, floods, and volcanism)

result in stand initiation and stem exclusion stages consisting

solely of even-aged cohorts of shade-intolerant tree species

(typically Nothofagus). Shade-tolerant tree species and tall

understory Chusquea bamboos typically do not establish in

abundance until the understory reinitiation stage. As the

initial cohort of shade-intolerant species becomes senescent,

dies and creates gaps, these gaps are preempted by the advance

regeneration of the shade-tolerant tree species and the

proliferation of bamboos. If the frequency and scale of

disturbance are sufficiently high, the shade-intolerant colonists

do not die before the next major disturbance and a true old-

growth stage (or steady-state stand) is not attained.

In areas of mixed-species rainforest where the stand-

initiating disturbance does not remove the understory of the

rainforest (e.g., a blowdown) the response is dominated by the

accelerated growth of plants that survive the disturbance rather

than by new establishment. In this case, it is mostly small

individuals of shade-tolerant trees and the Chusquea bamboos that

increase their growth rates, but shade-intolerant species such as

N. dombeyi may also regenerate in small numbers. In blowdowns of

mixed-species rainforest, scattered N. dombeyi may establish on

nurse logs and survive to maturity in low numbers, even with the

intense competition from bamboos and the released advance

39
regeneration of shade-tolerant species. In such stands, N.

dombeyi may or may not dominate the initial phases of stand

development depending on the abundance of suitable micro-sites

for seedling establishment and the understory composition of the

stand prior to disturbance.

The old-growth rainforests of the coastal mountains of

Chiloé and the Chilean Lake District are characterized by less

coarse-scale geologic disturbance because of their non-volcanic

lithology and fewer landslides. In these more stable habitats,

forest dynamics are more controlled by the occurrence of small

treefall gaps (Veblen et al. 1981, Veblen 1985b, Armesto and

Figueroa 1987, Armesto and Fuentes 1988). The most shade-

tolerant tree species are able to regenerate beneath a continuous

canopy cover, but their growth rates are enhanced by the creation

of small canopy gaps (Veblen 1985b, Armesto and Fuentes 1988).

Semi-shade-tolerant species, both subcanopy and main canopy in

potential stature, are entirelyy dependent on treefall gaps for

successful regeneration (Armesto and Fuentes 1988). Preemption

of canopy gaps by rapidly growing Chusquea bamboos can inhibit

tree regeneration, even of the most shade-tolerant tree species,

and slow the rate of gap filling. Tree species with some mode of

vegetative reproduction (e.g., root suckering, basal sprouting)

have an advantage in filling gaps (Veblen et al. 1981). In

landscapes that are characterized by lower rates of coarse-scale

disturbance, forests exhibit greater dominance by shade-tolerant

trees and appear to be closer to steady-state compositions

40
(Veblen et al. 1981, Armesto and Figueroa 1987).

Coarse-Scale Natural Disturbances

Disturbances of Tectonic Origin

Situated at a boundary between two major crustal plates, the

southern Andean region is an area of great volcanic and tectonic

activity. Ecological disturbances of geologic origin play a

major role in the dynamics of South American Nothofagus forests.

With the possible exception of New Guinea (Garwood et al. 1979),

no where else has tectonic disturbance been demonstrated to be as

important in forest dynamics (Veblen and Ashton 1978, Veblen et

al. 1992). On the wet western side of the Andes, in the

temperate rainforests of Chile, strong earthquakes such as the

1960 Valdivian earthquake measuring 8.5 on the Richter scale,

periodically, in a single event, trigger thousands of mass

movements (e.g. landslides, rockfalls, debris flows, and

mudflows) in the Andes. In addition to mass movements triggered

by earthquakes, many other mass movements are associated with

prolonged and torrential rainfall in this region of glacially

oversteepened slopes (Lara 1991).

Bare sites created by mass movements are favorable to the

establishment of the shade-intolerant Nothofagus species that

otherwise tend to be replaced by the numerous shade-tolerant tree

species of the Chilean rainforests (Veblen and Ashton 1978,

Veblen et al. 1980, Veblen et al. 1981, Lara 1991). Seedlings

and mature trees of Nothofagus spp. also are frequently observed

41
growing on recent volcanic deposits. Where buried by volcanic

ash deposits N. antarctica survives by sprouting adventititous

roots on branches and layering or by sprouting from the base

(Veblen et al. 1977). Following burial by a meter or more of

volcanic ash, N. pumilio has the capacity to develop a new

lateral root system closer to the new soil surface (Veblen et al.

1977). The rapid colonization and growth of Nothofagus on bare

surfaces is probably facilitated by their ectomycorrhizae and the

presence of nitrogen fixers (e.g. root nodule bacteria on

Discaria and Coriaria and blue-green algae on Gunnera). In areas

of high precipitation, the giant herb Gunnera chilensis which

supports nitrogen-fixing blue-green algae (Nostoc spp.) often

initially dominate bare sites exposed by landslides and are

replaced by Nothofagus within 20 to 30 years dense stands develop

(Fig. 10.11).

In contrast to the thousands of landslides on the Chilean

side of the Andes triggered by the 1960 earthquake, on the

Argentinean side of the Andes relatively few landslides were

triggered. However, intense shaking caused tree mortality and

affected tree growth rates (Veblen et al. 1992a). In some stands

located on unstable debris fans, most Nothofagus dombeyi and

Austrocedrus chilensis canopy trees died in 1960, allowing both

new establishment and accelerated growth of smaller surviving

trees of both species (Fig. 10.12; Veblen et al. 1992a).

Possible causes of death are damage to roots and probably a

sudden decline in soil moisture availability due to intense

42
shaking of the coarse substrate. Even in stands that show no

obvious signs of massive disturbance in 1960, dendroecological

analyses reveal sustained increases in growth releases,

apparently due to an earthquake-induced increase in treefall

frequency (Veblen et al. 1992a). On Tierra del Fuego, increased

treefall rates in N. pumilio were also associated with a strong

earthquake in 1949 centered near Lago Fagnano (Rebertus and

Veblen 1993b).

Since the early Spanish colonial period, high-magnitude

earthquakes have affected south-central Chile in 1575, 1737,

1751, 1837, 1939, and 1960 (Lomnitz 1970, Kanamori 1988) and

might also have affected Patagonian forests on the eastern slopes

of the Andes. In several stands of N. dombeyi and Austrocedrus

chilensis near Lago Traful in Argentina, A. chilensis tree-ring

chronologies indicate that the 1737, 1751 and 1837 earthquakes

affected tree growth similarly as the 1960 earthquake (Fig.

10.13). The chronologies show extremely narrow ring widths

coinciding with the earthquakes. Following one to a few years of

slow growth, the surviving trees accelerate their growth due to

decreased competition resulting from the death of other canopy

trees. The stronger release in the early 1960s probably reflects

the coincidence of the earthquake with two years of drought which

resulted in greater tree mortality and hence greater release

(Kitzberger et al. unpublished b).

Throughout the coastal area of southern Chile as well as in

the Andes, past earthquakes have caused extensive tree mortality.

43
For example, on the islands and the coast south of Chiloé Island,

stands of dead-standing trees lacking evidence of fire were

attributed to effects of tidal waves and tectonic subsidence

associated with the 1837 earthquake (Musters 1871, Rothkugel

1916). In areas near Lago Mascardi and Rio Puelo on the eastern

side of the Andes, extensive stands of dead-standing trees, in

areas apparently not burned, were observed by nineteenth century

explorers and attributed to raised water tables (Rothkugel 1916).

These changes in water table levels may have been due to local

mass wasting and glacier movements as suggested by Rothkugel

(1916) or they may have been the result of regionally extensive

tectonic events. Inundated coastal forests occur throughout the

southern Chilean Archipelago from just south of Chiloé Island to

Tierra del Fuego. On Tierra del Fuego in the area surrounding

Lago Fagnano extensive stands of Nothofagus were flooded and

killed when the area subsided in a 1949 earthquake (Goodall

1979).

Disturbances Associated with Glacial Processes

Establishment of Nothofagus spp. on recent glacial deposits

has been widely documented in Patagonia. In northern Patagonia

on glacier-covered Mt. Tronador (41o10'S), N. dombeyi establishes

in abundance on glacial deposits at low elevations in moist

habitats shortly after the deposits are exposed (Lawrence and

Lawrence 1959, Veblen et al. 1989a). Even-aged N. dombeyi stands

often develop on moraines that have become stabilized. N.

44
dombeyi is even capable of establishing and forming dense stands

on in-transit moraines (i.e., supraglacial debris) where mass

movements have deposited debris on top of the glacial ice (Veblen

et al. 1989a). At higher elevations and on drier substrates N.

antarctica may dominate the colonization of moraines and outwash

deposits.

At San Rafael Glacier (46o40'S), Heusser (1964) and Lawrence

and Lawrence (1959) described the establishment of Nothofagus

nitida, N. antarctica, and N. betuloides on moraines and outwash.

In far southern Patagonia establishment of N. betuloides, N.

pumilio and N. antarctica on moraines and outwash have been

reported for numerous glacial valleys (Pisano and Dimitri 1973,

Pisano 1978, Armesto et al. 1992). On some moraines near Tyndall

Glacier in Torres del Paine National Park (53oS), N. betuloides

appears to be the initial colonist and N. antarctica subsequently

invades as the moraine ages (Armesto et al. 1992). Armesto et

al. (1992) hypothesized that differential colonizing abilities of

Nothofagus species may be related to differential tolerances to

low nitrogen levels in skeletal glacial soils.

Other disturbance phenomena associated with glacial

processes in Patagonia include extensive flooding. For example,

periodic blockage of one arm of Lago Argentino (50o30'S) by the

Moreno Glacier results in a rise of water level of 8 to 15 m

(Pisano and Dimitri 1973). Partial melting of the ice block

periodically results in catastrophic flooding and destruction of

hundreds of hectares of Nothofagus forest.

45
 Blowdown

Patagonia is well known for its strong winds, and blowdowns

are an important disturbance in Patagonian Nothofagus forests

(Mutarelli and Orfila 1973, Eskuche 1973). The apparently even-

aged population structures common to many N. pumilio forests

appear to often result from large blowdowns. Beneath an even-

aged cohort there may be little or no successful regeneration of

N. pumilio until the stand is 200 or more years old. Since these

cohorts can cover areas of several hectares, this often gives the

impression of a non-regenerating forest, but where either the

senescence and death of large canopy trees occurs or the stand is

disturbed by exogenous disturbance (most commonly strong winds),

regeneration occurs (Alvarez and Grosse 1978, Schlegel et al.

1979, Schmidt and Urzua 1982, Armesto et al. 1992, Rebertus,

Veblen and Kitzberger unpublished data). Despite the prevalence

of the catastrophic mode for the regeneration of N. pumilio,

seedling establishment in large openings is sometimes inadequate

probably due to desiccation by exposure to strong winds (Roig et

al. 1985). Regeneration is frequently impeded by browsing by

livestock or introduced rabbits or hares (Mutarelli and Orfila

1973, Roig et al. 1985). Browsing by guanacos can nearly totally

block regeneration in small blowdowns, whereas in larger

blowdowns the severe browsing is limited to the perimeter of the

opening (Rebertus, Veblen and Kitzberger unpublished data).

Windstorms on Tierra del Fuego, causing the windthrow of

individual trees as well as the blowdown of entire stands, are

46
particularly important in controlling the structure of Nothofagus

forests (Puigdefábregas et al. in press, Rebertus and Veblen

1993b). Under certain circumstances of wind exposure,

topographic position, and substrate, gradual gap enlargement

results in a wave-like pattern similar to well documented fir

waves in the northeastern United States (Puigdefábregas et al. in

press, Rebertus and Veblen 1993a). Well-developed mortality

waves in N. betuloides forest occur at right angles to the coast

along the Beagle Channel and on Isla de los Estados, where strong

coastal winds and possibly salt spray may be the driving forces

(Kuhneman 1976). Further inland, waves do occur, but they are

less conspicuous and more irregular in shape and movement

(Rebertus and Veblen 1993a). Several inland mortality waves that

have been sampled indicate expansion rates of approximately 1

m/yr (for > 150 yrs), although these waves appear to expand

episodically in response to southerly windstorms rather than

gradually in response to prevailing winds (Rebertus and Veblen

1993a).

North of Lago Fagnano in the Sierra de las Pinturas of

Tierra del Fuego, large-scale blowdowns, sometimes involving > 1

km2, are the most conspicuous feature in the landscape. Indeed,

some areas are a complete mosaic of blowdowns of different ages.

The average return interval for blowdowns on exposed sites is c.

150-170 years (Rebertus, Veblen and Kitzberger unpublished data).

A shallow soil (10-30 cm) over fractured bedrock of sandstone and

conglomerate predisposes stands to blowdowns, especially even-

47
aged stands that have reached c. 25 cm median dbh. Extensive

blowdowns have occurred roughly every 20 years in this region,

usually from windstorms from the south (Rebertus, Veblen and

Kitzberger unpublished data). Valleys parallel to the wind and

upper leeward slopes are most vulnerable to blowdowns.

Snow Avalanches

On a more localized scale, disturbance by snow avalanches

can be important in Nothofagus forests (Eskuche 1973). In

subalpine forests of N. pumilio, infrequent but stand-replacing

snow avalanches typically result in the development of even-aged

stands (Veblen 1979, Veblen et al. 1981). Where avalanches occur

with greater frequency, however, they may prevent N. pumilio from

successionally replacing N. antarctica. Although N. pumilio is

longer lived and appears to be a better competitor, N.

antarctica's typically more prostrate, shrubby form and effective

vegetative reproduction through adventitious roots on damaged

branches, give it an advantage on sites frequently disturbed by

avalanches (Veblen et al. 1977). On steep windward slopes at

high latitudes, such as on Tierra del Fuego, avalanche paths are

often > 300 m long (Fig. 10.14).

Fire

In general, lightning is not common in southern Chile and

southwestern Argentina. It is extremely rare on the western side

of the Andes, but on the eastern side convective storms are more

48
common and lightning is an important source of fires. For

example, in the Argentinean Andes for the period 1938-1982

between c. 39 and 43o30'S, National Park authorities recorded

that > 15% of fires of known origin were started by lightning

(Bruno and Martín 1982). In southern Patagonia, Heusser (1987)

reports that all fires are either set by humans or ignited by

volcanism since lightning does not occur at these latitudes.

Nevertheless, lightning storms infrequently occur on Tierra del

Fuego (personal observation, A. Rebertus). Human-set fires, as

discussed below, have been significant in many types of

Nothofagus forests for several thousand years.

The abundance of fire-adapted species suggest that fire has

long been an important component of many Nothofagus ecosystems,

at least those occurring in drier habitats. For example,

Araucaria araucana, which grows in association with several

Nothofagus spp. in relatively dry habitats, is superbly adapted

to fire by its thick bark, protected terminal buds, and ability

to resprout after being burned (Veblen 1982b, Burns 1993).

Similarly, along the ecotone between Nothofagus forests and the

Patagonian steppe, most trees and shrubs (most notably Nothofagus

antarctica, Schinus patagonicus, and Lomatia hirsuta) vigorously

resprout after being burned. Likewise, in the northern part of

the Valdivian rainforest N. obliqua and N. alpina both resprout

after being burned.

Charcoal particles in sedimentary samples from c. 39oS in

Argentina to Tierra del Fuego, indicate that fire has been an

49
important disturbance in the zone of cool temperate Nothofagus

forests for at least several thousand years (Heusser 1987,

Heusser et al. 1988). Although more charcoal is found in

association with pollen assemblages indicating xeric, open

vegetation, the joint occurrence of charcoal with forest taxa

imply some incidence of pre-historic fire in forests. Fires were

set by Indians for purposes of communication, cooking, heating,

and hunting. In northern Patagonia early explorers describe

Indians using fires to hunt rheas and guanacos in the steppe

adjacent to forests (Cox 1863, Musters 1871, and Fonck 1900).

They also describe vast areas of burnt forest in the eighteenth

and nineteenth centuries that was attributed to Indian-set fires

(Cox 1863, Fonck 1900). Similarly, in the Chilean Lake District

the Mapuches (or Araucanians), well known for their warlike

habits, set fire to forests as both a defensive and aggressive

tactic (Encina 1954).

Within the Valdivian rainforest areas, intense fires promote

sprouting of N. obliqua and N. alpina, so large areas that were

cut and burned during the 1930s to 1950s are now dominated by

second-growth stands of these species. N. dombeyi is also a

common dominant of post-fire stands, but Chusquea bamboos sprout

vigorously after being burned and often dominate post-fire stands

and impede the establishment of all tree species. Where

Nothofagus occurs in pure stands in Argentina and far southern

Chile, all Nothofagus species develop dense even-aged stands

after fires if conditions are favorable (Fig. 10.15; Pisano and

50
Dimitri 1973, Roig et al. 1985, Veblen and Lorenz 1987,

Kitzberger and Veblen unpublished). The exceptions are for sites

lacking nearby seed sources, subject to severe browsing, exposed

to strong winds, on xeric slopes at high elevations, and/or where

soils are shallow.

The failure of regeneration after fire is most conspicuous

for N. pumilio forests in northern Patagonian Argentina on steep,

north-facing (xeric) slopes at high elevations, where we have

observed numerous burns > 40 years old lacking tree seedlings.

Given the lack of livestock at these sites and the proximity of

seed sources, the lack of tree regeneration may be due to fire-

induced edaphic changes or simply to opportunistic prior

establishment of a dense cover of herbaceous plants. Similarly,

in southern Patagonian Chile extensive burning of N. pumilio

forests typically results in only a narrow band (5 to 20-meters

wide) of dense tree establishment under the partial protection of

the forest margin. Lack of seedlings more distant from the

forest margin may be explained by a combination of poor seed

dispersal, herbivory by introduced hares and livestock, and/or

strong desiccating winds.

In the Valdivian and Patagonian rainforest regions, fires

typically shift the dominance from shade-tolerant species such as

Laurelia, Dasyphyllum, Saxegothaea, Aextoxicon, and Myrtaceae

towards the shade-intolerant Nothofagus. In the pure Patagonian

or Subantarctic Nothofagus forests, regeneration is typically to

the same species that dominated before the fire. An exception

51
occurs in locally unfavorable habitats in northern Patagonia,

where N. antarctica initially dominates after a fire but

eventually is replaced by the longer-lived N. dombeyi which

establishes jointly with N. antarctica (Veblen and Lorenz 1987).

More frequent burning generally favors dominance by N. antarctica

and numerous tall xerophytic shrubs which vigorously resprout

following fire.

Dieback Phenomena: Interactions of Disturbance, Climate and Stand

Dynamics

Abundant dead-standing trees and partial crown mortality are

conspicuous features of pure Nothofagus forests (Fig. 10.16).

The decline-disease theory of Manion (1981) emphasizes the

complexity of causal factors of dieback which can be classified

into (1) predisposing, (2) triggering or precipitating, and (3)

accelerating factors. Mueller-Dombois (1987, 1988), in his

theory of cohort senescence, has stressed the importance of even-

aged population structure leading to synchronized senescence and

mortality. Many South American Nothofagus species regenerate

following a catastrophic regeneration mode (sensu Veblen 1992)

and form even-aged stands. Thus, their demographic structure may

be an important predisposing condition for dieback. Occurrence

on suboptimal sites of either too little or too much moisture

availability may also be an important predisposing condition to

dieback. For example, Nothofagus dieback is particularly common

near their eastern limits where moisture availability is limited,

52
at high latitude sites characterized by strong winds, and at

sites of waterlogged soils.

On Tierra del Fuego in some stands of N. betuloides the

amount of dead-standing basal area is equal to the basal area of

live trees (Rebertus and Veblen 1993a). In many cases, these

appear to be old even-aged stands which are senescing and occur

at a range of spatial scales from fine- to coarse-scale patches.

No data are available on the triggering mechanisms of this

dieback. Once the canopy of a stand becomes more open due to the

initiation of dieback, strong winds on Tierra del Fuego are

undoubtedly an important accelerating factor in the spread of

dieback. Another form of dieback of N. betuloides on Tierra del

Fuego involves discrete patches of mortality over a few hectares

and is clearly associated with boggy conditions (Rebertus et al.

1993).

The previously described patches of dead-standing N. dombeyi

in northern Patagonia killed by the effects of the 1960

earthquake may also be interpreted within the framework of a

stand-level dieback. These are typically old, post-fire, even-

aged populations that are probably more susceptible to the damage

associated with earthquakes than would be younger trees.

Preliminary dendroecological data indicate that the occurrence of

earthquakes during periods of drought results in greater tree

mortality (Kitzberger, Veblen and Villalba unpublished b).

Substrate instability represents an important site factor

predisposing the stand to suffer from recurrent triggering

53
factors such as seismic vibrations and extreme drought. In some

cases, such as the 1960 mortality event, the earthquake event may

also be considered an accelerating factor where decline was

already initiated by the 1956-1962 drought. Dead-standing N.

dombeyi are also common in forests known to have been heavily

affected by volcanic ash deposition, such as the 1960 eruption of

Puyehue Volcano in Chile. However, the possible role of volcanic

ash deposition as an agent of mortality in these forests has not

been investigated.

Partial crown dieback is characteristic of N. antartica and

N. pumilio in the eastern parts of their distributions from

northern Patagonia to Tierra del Fuego and was formerly

attributed to recent climate change (Auer 1939, Kalela 1941).

Although the etiologies of these diebacks are not fully

understood, in the case of N. pumilio, insect damage, perhaps

associated with unusual weather may be the triggering mechanism.

During an outbreak of the caterpillar Ormiscodes cinnamomea

(Gentili and Gentili 1988) that we observed in northern Patagonia

in 1986, many hundreds of square kilometers of Nothofagus forests

in Nahuel Huapi and Lanín National Parks were temporarily

defoliated. This outbreak followed an exceptionally dry and warm

late winter and spring. Mild winters may favor survival of

insect larvae, whereas dry springs may make the hosts less

resistant to attack. The most severely affected species was N.

pumilio, but N. antarctica and N. dombeyi were also attacked in

1986 and in earlier outbreaks (Hosseus 1915). Our revisits to

54
attacked stands have indicated that the outbreak resulted in

relatively little tree mortality, but incomplete recovery has led

to an increase in trees with partial crown dieback. Our

preliminary survey of the degree of crown dieback in relation to

tree age indicated that trees classified as showing heavy dieback

were significantly older than trees showing slight dieback.

Attacks on Nothofagus by defoliators have also been reported for

Tierra del Fuego (Alfonso 1940) and adjacent mainland southern

Patagonia. Geometridae are important defoliators of N. pumilio

and may be reponsible for conspicuous defoliated patches of trees

in the Punta Arenas area (Welch 1988). Periodic insect

defoliation followed by incomplete recovery may contribute to the

abundance of N. pumilio with partially dead crowns throughout

Patagonia.

N. pumilio with partial crown dieback sometimes exhibit

symptoms of attack by wood boring insects (e.g. holes in the

bark, galleries in the phloem, and large scars surrounded by

callus tissue). Attacks of N. pumilio by wood borers

(Chilecomadia sp.) have been observed in Patagonia and Tierra del

Fuego (Petersen 1988, Gentili and Gentili 1988), and in the

stands we observed the attack was associated with severe crown

dieback.

Partial crown dieback in N. antarctica is also common along

the forest steppe ecotone throughout Patagonia and Tierra del

Fuego and may be related to cohort senescence. N. antarctica is

a relatively short-lived tree which rarely exceeds ages of 150

55
years, although occasionally trees over 200 years old can be

found. Most of the regeneration of N. antarctica occurs as basal

sprouts following the destruction of the main stem by fire

(Tortorelli 1956, McQueen 1976, Veblen and Lorenz 1988, Burns

1993). In the absence of fires, which rejuvenate stands by

stimulating vigorous sprouting, trees appear to senesce and

suffer from partial crown mortality (Veblen and Lorenz 1988).

Fire suppression since approximately 1920 over most of the range

of N. antarctica appears to have increased the proportion of

stands which are in senescent stages. For example, in the Nahuel

Huapi area mean fire return intervals near the ecotone with the

steppe are several-fold greater during the fire suppression

period (1930 to 1991) than during the pre-white settlement period

(pre-1898), when the native American population frequently burned

the ecotone for hunting purposes (Kitzberger et al. unpublished

a). Widespread burning with European settlement in the 1890s to

1920s followed by fire suppression has probably regionally

synchronized dieback by creating large areas of similarly aged N.

antarctica stands. Other factors, such as the semi-parasitic

Misodendrum spp. which is usually associated with dieback stems,

also may be involved. Howevever, there is a clear pattern of

young post-fire healthy stands versus abundant dieback in older

stands as well as greater ages of trees exhibiting more severe

dieback within the same stand (Veblen and Lorenz 1988).

Fine-scale Treefall Gap Dynamics

56
 Disturbance by fine-scale treefalls is important in all

Nothofagus forests, but has a fundamentally different influence

on stand development in mixed-species rainforests as opposed to

pure Nothofagus forests. In rainforests where Nothofagus spp.

occur with numerous shade-tolerant trees, small gaps are pre-

empted by the shade-tolerant trees and bamboos, so Nothofagus

spp. rarely establish in gaps (Fig. 10.17; Veblen 1985a, Armesto

and Figueroa 1987, Armesto and Fuentes 1988, Rebertus and Veblen

1993b). Thus, fine-scale treefalls accelerate succession towards

greater dominance by shade-tolerant tree species in the mixed-

species rainforests. In contrast, at higher elevations and

latitudes and on the drier side of the Andes, fine-scale

treefalls usually result in regeneration opportunities in these

pure or nearly pure Nothofagus forests (Veblen 1979, Veblen et

al. 1981, Schmidt and Urzua 1982, Veblen 1989b, Burns 1991,

Armesto et al. 1992, Innes 1992, Rebertus and Veblen 1993b,

Gutiérrez et al. in press). For example, on the drier eastern

side of the Andes at c. 40oS, N. dombeyi occurs in association

with the xerophytic conifer Austrocedrus chilensis and

regenerates in small treefall gaps (Veblen 1989a). Thus, the

regeneration modes of Nothofagus spp. vary according to the tree

and understory species with which they are associated (Veblen

1989b).

The regeneration of N. pumilio following small (i.e., single

tree) treefalls has been documented in the Chilean Lake District

(Veblen et al. 1981), Argentinean Lake District (Rusch 1992,

57
Burns 1991), in Chilean Aysén (Schlegel et al. 1979), and on

Tierra del Fuego (Gutiérrez et al. in press, Rebertus and Veblen

1993b). In the subalpine forests of the Chilean Lake District,

N. pumilio and N. dombeyi appear to occupy somewhat different

regeneration niches that enable them to regenerate in spite of

the strong competition from bamboos. In the northern part of

their distribution (i.e., north of c. 43oS) in subalpine forests,

bamboos often inhibit but do not totally impede the regeneration

of N. pumilio and N. betuloides. Beneath treefall gaps the

height and density of the bamboo is diminished, due probably to

longer lasting snow and cooler temperatures beneath gaps, so that

small numbers of N. pumilio seedlings are able to grow through

the bamboo layer and into the tree canopy (Veblen et al. 1981).

Although N. pumilio seedlings establish and persist for several

years beneath the bamboo cover, poised to respond to the creation

of a canopy gap, N. betuloides establishes primarily on large

logs or tree butts above the level of intense competition from

bamboo (Veblen et al. 1981). Where these elevated seedling sites

coincide with a canopy gap, regeneration may be successful.

Farther south (i.e., at c. 41o S) but still in the range of

understory bamboos, N. pumilio occurs in association with the

evergreen N. betuloides. The evergreen species appears to be

slightly more shade tolerant and occurs in greater abundance

beneath small treefall gaps, especially as seedlings established

on logs (Veblen et al. 1977, Veblen 1979). Larger canopy

openings (e.g. avalanche tracks) which are characterized by lower

58
minimum temperatures, appear to favor N. pumilio regeneration.

Similarly, farther south in Patagonia and on Tierra del

Fuego where bamboo is absent, N. betuloides appears more shade

tolerant than N. pumilio and is more abundant beneath small

treefall gaps (Schlegel et al. 1979, Rebertus and Veblen 1993b).

N. betuloides can remain suppressed in the understory for > 100

years, peristing as advance regeneration until one or more gaps

allow it to ascend into the canopy. Small individuals of both

species, however, are released by the creation of canopy gaps,

and both grow into the main canopy.

Also, on Tierra del Fuego, at low elevations near the coast

N. betuloides occurs in association with the subcanopy tree

Drimys winteri. In these wet forests small Drimys dominate the

understories and appear to restrict the regeneration of N.

betuloides (Rebertus and Veblen 1993b; Gutiérrez et al. in

press). Even though small Drimys dramatically outnumber N.

betuloides beneath small canopy gaps, the occurrence of a few

small trees of the latter species in approximately half the gaps

suggests that a low rate of fine-scale gap phase regeneration

does occur (Rebertus and Veblen 1993b). The regeneration of N.

betuloides is much more successful following larger blowdowns (>

c. 1000 m2) in these coastal rainforests. Further inland, Drimys

disappears and in its absence, the regeneration of N. betuloides

following canopy disturbances is abundant.

The few studies of treefall gap dynamics in South American

Nothofagus permit only a preliminary description of treefall gap

59
parameters. The largest gaps are created by N. dombeyi which may

exceed 50 m in height (Table 10.5). Gaps are often enlarged when

adjacent trees are toppled in domino-like fashion. On Tierra del

Fuego, most gaps were also the result of multiple treefalls

(median of 4 in several forest types), but expansion was gradual

and often decades apart from the intial event (Rebertus and

Veblen 1993). The size distributions of gaps are similar in

South American Nothofagus forests in the three areas that have

been studied (Chilean Lake District, northern Patagonia, and

Tierra del Fuego; Table 10.5) and also similar to New Zealand

Nothofagus forests (Stewart et al. 1991).

Wind-snapped trees were the most common gapmakers in old-

growth Nothofagus forests studied by Veblen (1985a) in Chile and

Stewart et al. (1991) in New Zealand; however, uprooting was

reported to be more common in northern Patagonia (Veblen 1989b)

and on Tierra del Fuego (Rebertus and Veblen 1993b). In

comparison with conifer-dominated rainforests of the Pacific

Northwest and with temperate forests in general, South American

Nothofagus forests are characterized by larger percentages of

uprooted trees (Peterson and Pickett 1991, Veblen and Alaback in

press). Possibly this reflects a greater resistance of many

angiosperm trees to bole breakage, higher intensity and more

frequent storms at high latitudes in southern South America,

and/or edaphic differences. The extremely shallow soils of

Tierra del Fuego undoubtedly predispose trees to uprooting, but

the intertwining roots of adjacent trees may help stabilize trees

60
until heartrot weakens the roots or trunk (Alfonso 1940). The

preponderance of uprooting in many forest types probably reflects

the severity of windstorms (c.f. Peterson and Pickett 1991) and

shallow, unstable soils that are susceptible to slippage when

wet. On Tierra del Fuego, even N. pumilio and N. antarctica, two

species notoriously prone to stem heart rot, are more prone to

uprooting than windsnap. Roots in shallow, rocky soils (e.g.

Lago Fagnano) are also subject to compression breaks during

windstorms. The 94% uprooting observed in blowdowns on Tierra

del Fuego (Rebertus et al. unpublished data) is probably the

highest reported in the literature (Peterson and Pickett 1991).

A striking feature of gaps in Tierra del Fuego was the

occurrence of multiple gapmakers per gap, many of which were

involved in secondary expansion events. In Fuegian forests it

appears that leeward edges of gaps are sometimes much more

susceptible to mortality than the forest as a whole. Such

expansion is often manifested in wave-like gaps, which have

expanded progressively downwind for > 100 m over a period of 100-

150 years (Rebertus and Veblen, 1993a), and more organized waves

that occur in series, similar to those found in Abies in North

America and Japan (Puigdefábregas et al. in press, Rebertus and

Veblen 1993b).

Turnover times from treefall gaps (based on the proportion

of forest in gap and the time required for gap closure) in the

few Nothofagus forests studied in South America range from 150 to

over 500 years (Table 10.5). Similar disturbance rates have been

61
reported for Nothofagus forests New Zealand (Stewart et al.

1991). However, we agree with Stewart et al. (1991) that such

estimates often fail to take into account variation in rates of

gap closure in small gaps, which tend to be rapidly infilled from

lateral growth of gap-edge trees; and large gaps tend to be

filled more slowly from growth of seedlings, saplings, and

suppressed understory trees. The spatial variation in

disturbance rate is underscored by the two to three-fold

difference in turnover in two N. pumilio forests on Tierra del

Fuego that are only 40 km apart. Nevertheless, these turnover

times are relatively long compared to temperate deciduous forests

in North America (Runkle 1985, Runkle 1990) and tropical rain

forests (Brokaw 1985) but are consistent with the greater

longevity of most Nothofagus.

Distributions of gap ages and releases of trees suggest that

gap formation in South American Nothofagus forests is highly

episodic and may be related to regionally extensive windstorms,

earthquakes, and stand-level dieback (Veblen 1985, Veblen et al.

1992, Rebertus and Veblen 1993b). The episodic nature of

treefalls and the likelihood of coarse-scale disturbances (fire

and blowdowns) during the lifespan of Nothofagus means that the

concept of treefall gap turnover time may be of limited value.

NOTHOFAGUS FOREST ECOSYSTEMS

Ecosystem-oriented studies that provide data on productivity

and nutrient cycling rarely have been conducted in South American

62
Nothofagus forests. Consequently, the following review at times

must rely on surrogate indicators of ecosystem processes, and

must be regarded as tentative.

Standing Crop - Basal Areas

Using basal area as a surrogate for standing crop, some

general trends can be identified. In south-central Chile P.

Donoso et al. (in press) have shown that the basal area of

similarly aged second-growth stands of N. obliqua and N. alpina

increases towards higher latitude. This implies that the greater

moisture availability at higher latitudes in this zone of summer

drought is an important determinant of basal area. Similarly,

the moisture limitation on basal area development is reflected by

lower basal areas near the northern limits of Nothofagus-

dominated old-growth stands (Donoso 1993, P. Donoso et al. in

press).

Basal area also varies consistently with elevation. For

example, along an elevational gradient at 40oS in the Andes of

south-central Chile basal area of Nothofagus-dominated forests

increases from the lowlands to peak values at 700 to 900 m

elevation and declines towards higher elevation (Donoso 1993).

The mid-elevation peak followed by a decline in the subalpine

zone of N. pumilio forest reflects the combination of high

precipitation and mild temperatures at mid-elevations, where

large N. dombeyi and N. alpina dominate. The maximum basal areas

of approximately 160 m2/ha in old-growth N. dombeyi-N. alpina-

63
Laurelia philippiana forests in south-central Chile are similar

to those reported for Nothofagus-angiosperm and Nothofagus-

gymnosperm forests in New Zealand (Chapter 3). At high latitudes

(c. 50 to 54oS), N. betuloides-N. pumilio forests attain basal

areas as high as 124 m2 (Pisano and Dimitri 1973), and the basal

areas of pure N. pumilio forests vary from 48 to 103 m2 (Schmidt

and Urzúa 1982).

Standing Crop - Wood Volumes

Although measurements of standing crop in South American

Nothofagus forests are too rare to permit comparisons and

generalizations, wood volume, for which many more data exist, is

a useful approximation to standing crop. However, wood volume

can only be taken as a rough index of standing crop, because

small diameters (typically < 35 cm dbh) are not included. As

expected, wood volumes are highest in the Valdivian rainforest

district and lowest where Nothofagus pumilio forms pure stands at

high latitude or high elevation. In south-central Chile, low and

mid-elevation old-growth stands dominated by N. obliqua, N.

dombeyi, and/or N. alpina attain wood volumes of c. 1300 m3/ha

(Donoso 1981a). Stands with the highest volume are late

successional stages, which still have many large (c. 1 to 2 m

diameter) individuals of the shade-intolerant Nothofagus. In

such stands, the Nothofagus account for c. 70% of the total

volume. High latitude stands of old-growth N. pumilio at c. 50

to 54oS have volumes of 400 to 600 m3/ha (Schmidt and Urzúa

64
1982).

Standing Crop - Weight of Biomass

The only comprehensive data on forest biomass are for Tierra

del Fuego (Table 10.6). In three monotypic old-growth stands of

N. antarctica, N. pumilio, and N. betuloides, total biomasses

were 248, 498, and 867 mt/ha, respectively (Frangi and Richter in

press). The two deciduous stands had 12 to 13% of their

biomasses below ground, whereas the evergreen stand had c. 19% of

its biomass below ground. The proportions of biomass of fine and

thick ( > 10 cm) roots were similar for the two deciduous

species. However, the biomass of thick roots of N. betuloides

was three times greater than that for N. pumilio, which indicates

a substantially greater allocation of biomass to structures used

in the absorption of water and nutrients by the evergreen

species. In contrast, the two deciduous species allocate

relatively larger percentages of their biomasses to structures

necessary for photosynthesis (Frangi and Richter in press). High

amounts of woody debris accumulation on the forest floor were

also found in these three stands: 83.4, 62.6, and 60.1 mt/ha in

the N. antarctica, N. pumilio, and N. betuloides stands,

respectively.

For three stands each of N. pumilio and N. betuloides on

Tierra del Fuego, Gutiérrez et al. (in press) report aboveground

biomasses of 127 to 284 mt/ha and 247 to 468 mt/ha, respectively.

In general, these stands show declining standing crops with

65
increasing elevation, except for the mid-elevation stand of N.

betuloides which had a lower than expected standing crop probably

due to poor drainage (Gutiérrez et al. in press). The biomasses

of the forests studied on Tierra del Fuego are similar to values

reported for New Zealand Nothofagus forests (Chapter 3).

However, there are no data on total biomasses of Nothofagus

forests in south-central Chile which contain the tallest and

largest diameter trees of southern South America. In the Chilean

and Argentinean Lake Districts understory Chusquea bamboos have

considerable biomasses. Understory standing crops of Chusquea as

high as 10 to 13 mt/ha have been measured at mid- (c. 700 m) and

high-elevations (1100 m) in old-growth stands in the Chilean Lake

District (Veblen 1982a, Veblen et al. 1980). In a post-fire

Chusquea culeou thicket lacking any trees, the aboveground

biomass was 156 to 162 mt/ha (Veblen et al. 1980) which is within

the range of values reported for Nothofagus forests on Tierra del

Fuego. These bamboo species synchronously flower and die

gregariously over large areas (Veblen 1982a) which undoubtedly

results in a major flux in biomass as well as nutrients.

Leaf Area Indices

Leaf area indices (LAI), measured only on Tierra del Fuego,

are 2.1 (plane m2 per m2 land surface) for N. antarctica, 1.7 to

2.8 for N. pumilio, and 2.5 to 6.5 for N. betuloides (Gutiérrez

et al. 1991, Frangi and Richter in press). Despite the

considerable range of variation, the evergreen species generally

66
had the highest LAIs. Although these LAIs are lower than in New

Zealand forests where they range from 4 to 9 (Chapter 3), the

number of South American stands for which LAI has been measured

is too small to allow conclusions.

Annual Productivity

Data on annual productivity in South American Nothofagus

forests are extremely sparse. Again, the only comprehensive

studies of annual productivity have been conducted on Tierra del

Fuego in Argentina. Gutiérrez et al. (in press) report annual

aboveground net production of 3.5 to 8.9 mt/ha and 4.8 to 8.4

mt/ha for three stands each of N. pumilio and N. betuloides,

respectively. Relations of annual wood production to leaf

biomass indicate that N. betuloides has a lower wood production

per unit of photosynthetic surface than N. pumilio (Gutiérrez et

al. 1991).

Fine litter (leaves, twigs and reproductive structures)

production has been measured in Nothofagus forests on Tierra del

Fuego and in the Chilean Lake District (Table 10.7). In

monotypic stands of Nothofagus on Tierra del Fuego, annual fine

litterfall ranges from 2.0 to 3.7 tn/ha. In the Chilean Lake

District where the growing season is longer, warmer and wetter,

fine litter production is much higher, ranging from 5.0 to 7.4

mt/ha. At c. 40oS, the highest values of fine litter production

are 7. 1 to 7.4 mt/ha/yr for low elevation secondary forests;

mid- and high-elevation old-growth forests range from 5.0 to 5.6

67
mt/ha/yr (Table 10.7). These values are similar to the highest

values reported for New Zealand lowland Nothofagus forests (5.0

to 7.3 mt/ha/yr) and somewhat greater than the 3.0 to 4.9

mt/ha/yr reported for high-elevation New Zealand forests (Chapter

3). In the forests sampled in the Chilean Lake District,

Chusquea bamboos accounted for 1.3 to 12.3% of the fine litter

production.

Nutrient Cycling.

Only preliminary data on nutrient cycling are available for

South American Nothofagus forests, and these are limited to three

stands dominated by N. antarctica, N. pumilio, and N. betuloides

on Tierra del Fuego (Frangi and Richter in press). Total

nutrient content of the living biomass is much greater in N.

betuloides as expected due to its greater total biomass.

However, the concentration (i.e. weight of ash per weight of

biomass) also is greater for the evergreen species (1.4% versus

1.0 and 0.8% for N. pumilio and N. antarctica, respectively).

The evergreen stand stored 65% of the total ash below ground, in

contrast to 26% and 52% for N. antarctica and N. pumilio,

respectively.

Frangi and Richter (in press) found the nutrient content of

the coarse woody debris on the forest floor to be lower in the N.

betuloides stand. However, the fine litter (leaves and twigs) of

the N. betuloides stand had the highest concentrations for most

nutrients. The leaf decomposition rate was substantially slower

68
in the sclerophyllous evergreen species than in the two deciduous

species (17% versus 56 to 59% per year). Prior to leaf

abscission of N. pumilio, 40 to 60% of the critical nutrients (N,

K, and P) are translocated (Richter and Frangi 1992). The rates

of decomposition of large branches and trunks were estimated only

for N. pumilio and were found to be extremely slow. Large

branches require an estimated 53 years to decompose, and for a

trunk to lose 99% of its mass requires an estimated 500 years

(Frangi and Richter in press). Such slow rates of decomposition

are consistent with the cool temperatures of Tierra del Fuego.

At local scales, the distribution of N. pumilio and N.

betuloides at high latitudes is at least partially explained by

nutrient differences in the soils (Gutiérrez et al. 1991). The

greater waterlogging of soils beneath N. betuloides appears to

result in deficiencies of nutrients, especially phosphorus, in

comparison with soils beneath N. pumilio (Gutiérrez et al. 1991).

Due to the lack of oxygen in the more waterlogged soils beneath

N. betuloides, mycorrhizae are also reported to be much less

abundant than they are beneath N. pumilio. The evergreen

sclerophyllous leaf habit of N. betuloides may be advantageous

for the accumulation of nutrients when growth is limited by low

nutrient availability (Gutiérrez et al. 1991). At these

nutrient-poor sites, the greater longevity of leaves, maintenance

of leaves throughout the year, and translocation of nutrients

from abscissing leaves all may be adaptations to conserve

nutrients (Gutiérrez et al. 1991). Such a strategy is only

69
successful where minimum temperatures are high enough to permit

winter persistence of leaves.

IMPACTS OF MODERN HUMANS

Introduced Animals

European contact quickly changed the assemblage of large

browsing mammals affecting Patagonian forests. From early

colonial Spanish settlements on the Pacific coast of Chile and

the La Plata estuary of eastern Argentina, livestock escaped and

quickly spread to the Nothofagus region (Fonck 1900). For

example, horses, sheep and cattle were present by the late

eighteenth century in the Nahuel Huapi area of northern Patagonia

(Machoni 1732, Cox 1863, Fonck 1900). By the end of the

nineteenth century livestock raising was widespread in southern

Chile and southwestern Argentina (Martinic 1985, Roig et al.

1985). In general, the drier Nothofagus forests have been

subject to more livestock impact than the rainforests. Even in

many reserves and national parks, livestock were common until the

1970s and are still present in many.

Exotic deer, upon which a lucrative sport hunting industry

depends, have also attained high population densities and have

seriously affected Nothofagus forests (Daciuk 1978, Ramírez et

al. 1981, Ramilo 1985, Veblen et al. 1989b). Red deer (Cervus

elaphus), which were introduced early in the 20th century from

Europe, are the most widespread exotic deer (Godoy 1963, Daciuk

1978). Red deer are now common in northern Patagonia, especially

70
in the ecotone between the forest and steppe, and also occur in

the Valdivian rainforest region of Chile (Ramírez et al. 1981,

Ramilo 1985). Fallow deer (Dama dama) and axis deer (Axis axis)

were introduced to the Argentinean Lake District in the 1930s and

the former are now common on Isla Victoria in Nahuel Huapi Lake

(Daciuk 1978).

Both deer and livestock have had major influences on the

structure and composition of Nothofagus forests (Boelcke 1957,

Daciuk 1978, Ramírez et al. 1981, Martín et al. 1985, Veblen et

al. 1989b, Veblen et al. 1992b). The principal difference

between deer and livestock browsing is the greater selectivity of

the former. In general, Nothofagus spp. are palatable to both

deer and livestock, and heavy browsing pressure can seriously

impede tree regeneration (Fig. 10.18). Heavy browsing leads to

an increase in relative abundance of spiny shrubs (e.g. Berberis

spp.), an overall decrease in the size and abundance of woody

plants, an increase in the importance of the herbaceous stratum,

and an increase in the relative abundance of exotic plants

(Ramírez et al. 1981, Martin et al. 1985, Roig et al. 1985, Brión

et al. 1988, Veblen et al. 1989b, Veblen et al. 1992b).

The nature of animal impacts in Nothofagus forests depends

on the stage of stand development. In post-fire stands in

northern Patagonia, for example, browsing mammals significantly

reduce height growth rates of N. dombeyi (and associated tree

species) in the stem-initiation phase of stand development and

generally retard the development of a complete tree cover. Once

71
stands have become taller and denser in the stem-exclusion stage

(sensu Oliver 1981), browsing is unimportant because there are no

small shade-tolerant trees. As post-fire stands develop into the

understory-reinitiation and old-growth stages (sensu Oliver 1981)

and treefalls create canopy openings large enough for the

establishment of seedlings of the shade-intolerant tree species,

browsing impacts again become critical (Veblen et al. 1989b,

Veblen et al. 1992b).

Introduced European hares (Lepus capensis) and wild boars

(Sus scrofa) have colonized nearly all suitable habitats of

mainland Patagonia (Daciuk 1978). The rooting activities of wild

boars locally alter grasslands and forest openings, especially in

the Andes of Rio Negro and Chubut provinces, Argentina (Daciuk

1978). European hares occur throughout continental Patagonia and

often attain high population densities, especially in open

habitats. They commonly damage Nothofagus seedlings,

particularly during winter when tall seedlings are exposed above

the snow and little alternative forage is available. Although

hares and boars do not occur on Tierra del Fuego, introduced

rabbits (Oryctolagus cuniculus) and beavers (Castor canadensis)

are severe pests there (Goodall 1979). In some N. pumilio stands

on Tierra del Fuego, rabbits prevent tree regeneration beneath

canopy gaps (Rebertus and Veblen 1993b). Beavers were introduced

to Tierra del Fuego in 1945-1946 to permit the establishment of a

fur industry, which was never developed (Lizarralde et al. 1990).

They have since spread throughout the Magellanic Archipelago and

72
threaten to invade mainland Patagonia. Beavers cut and kill many

N. pumilio, N. betuloides, and N. antarctica, from rainforest to

steppe and from sea level to above the upper forest limit.

Flooding associated with beaver dams presents a far greater

danger to Nothofagus forests than cutting, particularly in low,

boggy floodplains (Fig. 10.19).

Fire and Forest Clearance

In the Chilean Lake District, massive forest clearing and

burning began in the mid-nineteenth century with the arrival of

German colonists (Perez 1958). Nearly all the low elevation

forests were converted to agriculture in one of the most massive

and rapid deforestations in the history of Latin America. Vast

areas of forests with N. obliqua in the Central Depression were

converted to agricultural use (Donoso 1983). During the late

nineteenth and early twentieth centuries, extensive areas of mid-

elevation Nothofagus forests were burned in the Chilean and

Argentinean Lake Districts.

In northern Patagonia, the Indian war of 1879-83 drastically

reduced the native American population and was soon followed by a

period of deliberate forest burning and clearing for pasture

creation (Moreno 1897, Willis 1914). The summer of 1913-14,

which followed several years of drought, was a time of

exceptionally severe forest fires (Willis 1914, Rothkugel 1916).

Frequent burning from c. 1884 to 1920 is reflected in the

abundance of charcoal near the surface of sedimentary records

73
(Markgraf 1983), and by the abundance of 70 to 100 year-old even-

aged Nothofagus forests in northern Patagonia (Veblen and Lorenz

1987, Veblen and Markgraf 1988). In 1915, 25% of the forested

area of Argentinean Patagonia (including Tierra del Fuego) had

been recently burned (Rothkugel 1916). In Nahuel Huapi National

Park the incidence of fire in areas of mesic Nothofagus forest

increased several-fold during the early 1900s compared to the

pre-European period (Veblen et al. 1992a).

In southern Patagonia, extensive forest burning was also

characteristic of white colonization in the late nineteenth and

early twentieth centuries. For example, in the Chilean Province

of Aysén, where colonization began only in the 1920s, vast areas

of Nothofagus forests were burned to extend the area of sheep

raising. Of the original five million hectares of forests in

Aysén, 50 percent has been destroyed during this century

(Corporación Nacional Forestal, Coyhaique Office, unpublished

data, 1978). White patches of standing dead trees are mute

testimony to the extent of burning from the 1920s to 1940s and to

the slow rate of forest recovery in Aysén. Severe accelerated

erosion often followed burning. Further west, in the Chonos and

Guaitecas Archipelagoes and on the Taitao Peninsula, extensive

areas of forest were intentionally burned to facilitate logging

of the fire-resistant timber of Pilgerodendron uviferum (ciprés

de las Guaitecas) (Grosse 1979).

Forest Conservation and Management

74
 The Nothofagus forests of southern Chile and Argentina have

been subject to mostly destructive exploitation rather than

effective management, and to a substantial extent they have been

converted to plantations of exotic trees. Since the 1950s, the

area planted annually to Pinus radiata in Chile has grown

exponentially and in 1992 amounted to 1.55 million ha (Cerda et

al. 1992). Since the 1970s incentives by the Chilean government

to promote reforestation have greatly aided the development of

exotic plantations, but they have also permitted the conversion

of some native forest to exotic plantation (Lara and Veblen

1992). This has been particularly severe in south-central Chile.

Similarly, exotic pines have been extensively planted in

Argentina along the ecotone between Nothofagus forests and

Patagonian steppe. In both countries, there is much more

silviculture of exotic plantations than of the native forest.

In Chile, since 1986 large-scale production of wood chips

from the native forest has had a severe impact on the extent and

quality of the native forest resource. During the late 1980s to

early 1990s, an estimated 13,200 ha/yr of native forest were

being converted to other uses, and the quality and productive

capacity of another 108,600 ha/yr were being seriously reduced.

In contrast, only 2000 to 5700 ha of native forest were subject

to any type of silvicultural management in Chile (Ormazábal

1992). In Argentina, conversion of Nothofagus forest to

plantations of exotic trees has not been extensive and the recent

rate of forest degradation through destructive logging practices

75
has not been high. However, significant areas of Nothofagus

forest were destructively logged and/or burned during the first

half of the twentieth century. In disturbed areas, exotic

species are typically abundant and some aggressive invaders such

as Rosa eglanteria are locally dominant (Brion et al. 1988).

Since the mid-1970s there has been substantial silvicultural

research conducted in South American Nothofagus forests but

relatively little of the resulting knowledge has been put to use

in forest management (Bava and Schmaltz 1992, Donoso and Lara in

press). The minimal management of native forests in Chile and

Argentina is unfortunate given the potential inherent in the

timber quality and rapid growth of these forests. The

traditional form of forest exploitation in Nothofagus forests in

southern Chile and Argentina has consisted of high grading in

which the best-formed individuals of the most valuable timber

species are extracted. This has degraded the timber quality of

the resource by removing the more commercially valuable species

as well as the silviculturally superior genetic material. In

mixed-species rainforests, selective logging has also favored the

regeneration of the shade-tolerant tree species at the expense of

the more valuable Nothofagus. The relatively small canopy

openings created are filled by the advance regeneration of the

shade-tolerant tree species and Chusquea bamboos (Veblen 1982a).

In the mixed-species forests of the Chilean and Argentinean Lake

Districts, the best regeneration of Nothofagus has occurred where

the silvicultural intervention was more intensive, such as after

76
the cutting of patches of several hectares. Where small

clearcuts, often burned, are not subject to overly intense use by

livestock, dense secondary stands of N. obliqua, N. alpina, and

N. dombeyi typically develop. The former two species coppice and

all three species form dense and often nearly pure secondary

stands which are relatively easily managed by thinnings.

The silvicultural management of secondary stands of N.

dombeyi, N. obliqua, and N. alpina is relatively simple and

effective (Gutiérrez 1989). For example, thinnings of N. alpina

stands at age 10 to 15 years can produce 247 m3/ha of sawtimber

from the thinning plus a total of c. 690 m3/ha in the final cut

after 60 years (Grosse 1989). Although most management of the

native forest in the Chilean Lake District has consisted of

thinnings of post-fire secondary stands, effective silviculture

of some of the old-growth Nothofagus forests has also been

demonstrated. For example, in mid-elevation forests in the Andes

old-growth stands which were initially high graded have been

subjected to successive thinnings to transform them into

productive stands with adequate annual volumetric increments

(Donoso and Lara in press). Similarly, in Lanín National Reserve

in the Argentinean Lake District N. dombeyi-N. alpina forests are

now being subjected to silvicultural trials and forest management

plans (Luis Chauchard, pers. comm.).

In the Chilean provinces of Aysén and Magallanes, N.

pumilio, which produces excellent wood, has been extensively

managed. Thinning trials with this species indicate that 400 to

77
600 m3/ha of sawtimber can be produced in a 70 to 80-year

rotation (Vera 1985). Regeneration experiments in Argentinean N.

pumilio forests have also provided an initial basis for forest

management (e.g. Mutarelli and Orfila 1973), and currently there

is substantial silvicultural research underway in these forests

(Bava and Schmaltz 1992).

In both Chile and Argentina, large areas of Nothofagus

forest are protected in numerous national parks and reserves. In

particular, the inclusion of a large percentage of Nothofagus

forests in national parks and the low rate of forest exploitation

in Argentina are highly favorable for forest conservation.

However, the total area of Nothofagus forest in Argentina is

small compared to Chile. In both countries, although the number

of reserves and their surface areas are impressive (Ortiz et al.

1993), the protection of Nothofagus forests is not necessarily

adequate. It is unclear if the areas in parks and reserves

encompass a sufficiently large and diverse mosaic of environments

to adequately protect Nothofagus ecosystems. Regional- and

landscape-scale studies are needed for assessing the sufficiency

of present parks and reserves. For example, in the coastal

mountains of Chile most parks are small and their lack of

connection to larger Andean parks may be detrimental to the

survival of some species (Simonetti and Armesto 1991).

Similarly, it is not known whether individual parks are larger

than the "minimum dynamic area" (sensu Pickett and Thompson 1978)

which includes all successional stages of the present plant

78
communities. Furthermore, in many of the parks and reserves the

forests were burned and/or logged prior to the creation of the

reserve and there are serious invasions of the disturbed areas by

exotic species (Brión et al. 1988). In most parks and reserves

there are severe impacts from introduced animals such as

livestock, deer, and beavers. Some species and forest types are

not adequately represented in reserves. For example, unlogged

old-growth stands of N. alpina are not included in any Chilean

reserves and are represented by only a small area in a single

Argentinean national park. Information on the amount and

distribution of genetic diversity is also fundamental for

planning the conservation of Nothofagus species (e.g. Premoli in

press).

Global Change and South American Nothofagus Forests

Current general circulation models based on CO2 doubling

predict that southern South America will not experience warming

or moisture deficits as dramatic as comparable latitudes in the

Northern Hemisphere. Nevertheless, Manabe and Wetherald (1987)

indicate a 3 to 4oC increase in both summer and winter

temperature between 30 and 55oS in South America which would have

significant impacts on South American Nothofagus forests.

In South America, Nothofagus-dominated forests generally

occur at sites that are suboptimal for tree growth due to

climatic or edaphic factors. Thus, along a west-to-east trans-

Andean gradient at c. 40oS, the Valdivian rainforest of south-

79
central Chile is replaced by monotypic stands of Nothofagus,

which in turn give way to xeric woodlands and Patagonian steppe.

This is one of the world's steepest gradients from rainforests to

semi-arid steppe, and it is likely that it will be highly

sensitive to anthropogenic changes in temperature and

evapotranspiration. The paleoecological record indicates that

the occurrence of Nothofagus forests in ecotones between forest

and non-forest vegetation is associated with high sensitivity to

the effects of past climate fluctuation (Markgraf 1983, 1987).

Global warming is likely to have a major impact on

Nothofagus forests by changing disturbance regimes. For example,

the area burned per year in northern Patagonian mesic Nothofagus

forests is highly correlated with spring and summer water

deficits (Kitzberger et al. unpublished a). These drier

conditions are associated with a more intense and more southerly

location of the southeastern Pacific anticyclone which controls

the westerly flow of moist Pacific air into the continent. Thus,

relatively slight changes in atmospheric circulation patterns may

result in a dramatic increase in the occurrence of fire in South

American Nothofagus forests. Although most fires in these

forests are set by humans, droughts clearly predispose the

forests to more extensive burning. For example, in northern

Patagonia, the time of most extensive forest burning coincided

both with European settlement and prolonged drought from c. 1911

to 1917 (Veblen and Lorenz 1988, Veblen et al. 1992a).

Similarly, preliminary studies suggest linkages between weather

80
and insect outbreaks and Nothofagus stand dieback (Veblen,

Kitzberger and Villalba unpublished data). The influences of

climatic variation on disturbance regimes and forest dynamics in

South American Nothofagus forests is a major research challenge.

In addition to the potential effects of global warming on

South American Nothofagus forests, more direct effects of

atmospheric changes may be significant. For example, during late

September, 1992, stratospheric ozone depletion normally

restricted to Antarctica drifted over the southern tip of South

America as far north as 53oS (Waters et al. 1993), exposing

Nothofagus forests to high UV radiation. These events and links

between chloroflourcarbons and ozone depletion have kindled fears

that UV may harm many forms of life in this region (Kamm 1993).

In plants, high levels of UV radiation are known to depress

flower development, induce leaf abscission and loss of apical

dominance, and increase the root to shoot ratio (Kozlowski et al.

1991). Although leaves possess many protective features that

minimize damage from UV radiation, pollen is very susceptible to

injury or death from UV-B radiation. These possible influences

of increased UV radiation on Nothofagus have not yet been

investigated.

Despite the potential influences of global climatic change

on South American Nothofagus forests, many of these ecosystems

are relatively free from atmospheric depositions from urban and

and industrial sources. Thus, the monitoring and investigation

of these ecosystems are essential for the preservation and

81
management of them, and also can provide important comparative

data in relation to northern hemisphere temperate forests which

have been extensively affected by atmospheric deposition.

ACKNOWLEDGMENTS

For commenting on the manuscript we thank A. Premoli and R.

Villalba. Research support was provided by the National Science

Foundation, National Geographic Society and the Council on

Research and Creative Work of the the University of Colorado.

REFERENCES

Alberdi, M. 1987. Ecofisiología de especies chilenas del género

Nothofagus. Bosque 8: 77-84.

Alberdi, M., M. Romero, D. Rios, and H. Wenzel. 1985. Altitudinal

gradients of seasonal frost resistance in Nothofagus

communities of southern Chile. Acta Oecol. Plant. 6: 21-30.

Alfonso, J.L. 1940. Algunas consideraciones sobre los bosques de

Tierra del Fuego. Ingeniería Agronómica 1: 10-23.

Alvarez, S. and H. Grosse. 1978. Antecedentes generales y

análisis para el manejo de Lenga (N. pumilio) en Alto Mañihuales,

Aysén. Thesis, Universidad de Chile, Santiago, Chile.

Armesto, J.J., I. Casassa, and O. Dollenz. 1992. Age structure

and dynamics of Patagonian beech forests in Torres del Paine

National Park, Chile. Vegetatio 98: 13-22.

Armesto, J.J. and J. Figueroa. 1987. Stand structure and dynamics

82
 in the temperate rain forests of Chiloé Archipelago, Chile. J.

Biogeog. 14: 367-376.

Armesto, J.J. and E.R. Fuentes. 1988. Tree species regeneration

in a mid-elevation, temperate rain forest in Isla de Chiloé,

Chile. Vegetatio 74: 151-159.

Armesto, J.J. and R. Rozzi. 1989. Seed dispersal syndromes in the

rain forest of Chiloé: evidence for the importance of biotic

dispersal in a temperate rain forest. J. Biogeog. 16: 219-226.

Arroyo, M.T.K., M. Riveros, A. Peñaloza, L. Cavieres, and A.M.

Faggi. In press. History and regional richness patterns of the

cool temperate rainforest flora of southern South America. In

R.G. Lawford, P. Alaback, and E.R. Fuentes (eds.), High

Latitude Rain Forests and Associated Ecosystems of the Western

Coast of the Americas: Climate, Hydrology, Ecology and

Conservation. Springer-Verlag, New York.

Auer, V. 1939. Der Kampf zwischen Wald und Steppe auf Feuerland.

Petermanns Mitteilungen 6: 193-97.

Barros, V., Cordon, V., Moyano, C., Mendez, R., Forquera, J. and

Pizzio, O. 1983. Cartas de precipitación de la zona oeste

del las provincias de Rio Negro y Neuquén. Report Facultad

de Ciencias Agrarias Universidad Nacional del Comahue, Neuquén,

Argentina.

Bava J.O. and C.J. Puig. 1992. Regeneración natural de lenga.

Análisis de algunos factores involucrados. In J. Bava and J.

Schmalz (eds.), Actas Seminario de Manejo Forestal de la Lenga

y Aspectos Ecológicos Relacionados. CIEFAP, Publicación Técnica

83
 8. Esquel, pp. 85-110.

Bava, J. and J. Schmalz. (eds.) 1992. Actas Seminario de Manejo

Forestal de la Lenga y Aspectos Ecológicos Relacionados. CIEFAP,

Publicación Técnica 8. Esquel, Argentina.

Becker, J. 1981. Estudio de producción de litter en bosques

latifoliados del sur de Chile. Thesis, Universidad Austral de

Chile, Valdivia, Chile.

Boelcke, O. 1957. Comunidades herbáceas del norte de Patagonia y

sus relaciónes con la ganadería. Revista de Investigaciones

Agrícolas 11: 5-98.

Bridges, E.L. 1948. Uttermost Part of the Earth. Hodler and

Stoughton Ltd. London, England.

Brión, C., D. Grigera, J. Puntieri, and E. Rapoport. 1988. Plantas

exóticas en bosques de Nothofagus. Comparaciones preliminares

entre el norte de la Patagonia y Tierra del Fuego. Monografías

de la Acad. Nac. de Ciencias Exacatas, Físicas y Naturales 4: 37-

47.

Brokaw, N.V.L. 1985. Treefalls, regrowth, and community structure

in tropical forests. In S.T.A. Pickett and P.S. White (eds.),

The Ecology of Natural Disturbance and Patch Dynamics, Academic

Press, New York, pp. 53-71.

Bruno, J. and G. Martín. 1982. Los incendios forestales en los

Parques Nacionales. Unpublished report, Administración de

Parques Nacionales, Buenos Aires, Argentina.

Burns, B.R. 1991. Regeneration dynamics of Araucaria araucana.

Ph.D. thesis, University of Colorado, Boulder, Colorado, U.S.A.

84
-----. 1993. Fire-induced dynamics of Araucaria araucana-

Nothofagus antarctica forest in the southern Andes. J. Biogeog.

20: 669-685.

Burschel, P., C. Gallegos, O. Martínez, and W. Moll. 1976.

Composición y dinámica regenerativa de un bosque vírgen mixto

de raulí y coigüe. Bosque 2: 55-74.

Butin, H. 1969. Studien zur morphologie und biologie von

Mikronegeria fagi Diet & Neg. Phytopath. Z. 64: 242-257.

Carrillo, R. and L. Cerda. 1987. Zoofitófagos en Nothofagus

chilenos. Bosque 8: 99-103.

Cerda, I., J. Olavarría, and M.J. Abalos. 1992. El sector

forestal de Chile.- Logros y desafíos. Instituto Forestal -

Santiago, Chile.

Correa, A., J.J. Armesto, R. Schlatter, R. Rozzi, and P. Torre-

Murúa. 1990. La diete del Chucao (Scelorchilus rubecula), un

passeriforme terrícola endémico del bosque templado húmedo de

Sudamérica Austral. Rev. Chil. His. Nat. 63: 197-202.

Cox, G. 1863. Viajes a las regiones septentrionales de Patagonia

1862-1863. Anales de la Universidad de Chile 23: 3-239 and 437-

509.

Cwielong, P. and M. Rajchenberg. 1992. Hongos asociados a

pudriciones en madera de lenga. In J. Bava and J. Schmalz (eds.),

Actas Seminario de Manejo Forestal de la Lenga y A s p e c t o s

Ecológicos Relacionados. CIEFAP, Publicación Técnica 8. Esquel,

pp. 1-2.

Daciuk, J. 1978. Notas faunísticas y bioecológicas de Península

85
 Valdés y Patagonia: IV. Estado actual de las especies de

mamíferos introducidos en la subregión araucana (Rep. Argentina)

y del grado de coacción ejercido en el ecosistema. Anales de

Parques Nacionales 14: 105-130.

Delgado, C. 1986. Caracterización de un renoval de coigüe (N.

dombeyi) en el Sector Laguna P. Aguirre Cedra-Monte Picaflor,

Aysén. Thesis, Universidad de Concepción, Concepción.

Donoso, C. 1974. Dendrología: Arboles y Arbustos Chilenos.

Universidad de Chile, Facultad de Ciencias Forestales, Manual

No. 2. 142 pp.

-----. 1981. Los tipos forestales nativos de Chile. Documento

de trabajo n. 38. Publicación FAO-Chile.

-----. 1983. Modificaciones del paisaje forestal chileno a lo

largo de la historia. In Symposio Desarrollo y Prespectivas de

las Disciplinas Forestales.

Universidad Austral de Chile. Valdivia, pp. 365-438

-----. 1984. Hibridación natural entre Nothofagus betuloides

(Mirb.) Oerst. y N. nitida (Phil.) Krasser. Medio Ambiente 7:

9-16.

-----. 1987. Variación natural en especies de Nothofagus en

Chile. Bosque 8: 85-97.

-----. 1993. Bosques Templados de Chile y Argentina. Variación,

Estructura y Dinámica. Editorial Universitaria. Santiago,

Chile.

Donoso, C. and J. Atienza. 1983. Hibridación natural entre

especies de Nothofagus siempreverdes en Chile. Bosque 5: 21-34.

86
Donoso, C. and A. Cabello. 1978. Antecedentes fenológicos y de

germinación de la especies leñosas chilenas. Cs. Forestales 1:

31-42.

Donoso, C., M. Cortés, and B. Escobar. 1992. Técnicas de vivero y

plantaciones para Roble. Documento Técnico n. 62, Chile

Forestal.

Donoso, C., B. Escobar, and M. Cortés. 1991a. Técnicas de vivero

y plantaciones para Raulí. Documento Técnico n. 53, Chile

Forestal.

Donoso, C., B. Escobar, and M. Cortés. 1991b. Técnicas de vivero

y plantaciones para Coigüe. Documento Técnico n. 55, Chile

Forestal.

Donoso, C., B. Escobar, and J. Urrutia. 1985. Estructura y

estrategias regenerativas de un bosque virgen de Ulmo (Eucryphia

cordifolia Cav.) - Tepa (Laurelia philippiana Phil.) Looser en

Chiloé, Chile. Rev. Chil. His. Nat. 58: 171-186.

Donoso, C., R. Grez, B. Escobar, and P. Real. 1984. Estructura y

dinámica de bosques del tipo forestal siempreverde en un sector

de Chiloé Insular. Bosque 5: 82-104.

Donoso, C. and A. Lara. In press. Utilización del bosque nativo

en Chile: pasado, presente y futuro. In J. Armesto, M. Kalin

and C. Villagrán (eds.), Ecología del Bosque Nativo de Chile.

Donoso, C., J. Morales, and M. Romero. 1990. Hibridación natural

entre Roble (Nothofagus obliqua (Mirb.) Oerst.) y Raulí

(Nothofagus alpina (Poepp. y Endl.) Oerst.) en bosques del sur

de Chile. Revista Chilena de Historia Natural 63: 49-60.

87
Donoso, P. 1988. Caracterización y proposiciones silviculturales

para comunidades de Roble y Raulí en el área de protección

"Rodal 7 Tazas" VIII Región. Bosque 9: 103-114.

Donoso, P., C. Donoso, and V. Sandoval. In press. Caracterización

y crecimiento de renovales de Roble y Raulí en su distribución

latitudinal en Chile. Bosque 14.

Encina, F.A. 1954. Resumen de la Historia de Chile, Vol. 1. Zig-

Zag, Santiago.

Eskuche, U. 1973. Estudios fitosociológicos en el norte de

Patagonia. I. Investigación de algunos factores de ambiente en

comunidades de bosque y de chaparral. Phytocoenología 1: 164-

213.

Fonck, F. 1900. Viajes de Fray Francisco Menéndez a Nahuelhuapi.

C.F. Niemeyer, Valparaiso, Chile.

Frangi, J. L. and L.L. Richter. In press. Estructura y función de

ecosistemas forestales fueguinos: algunos resultados

preliminares del Proyecto Subantarctis. In J. Puigdefábregas,

F. García Novo, J. Frangi, and A. Vila (eds.), La Tierra del

Fuego: Los Sistemas Naturales y su Ocupación Humana. Consejo

Superior de Investigación Científica, Madrid, Spain.

Garwood, N.C., D.P. Janos, and N. Brokaw. 1979. Earthquake caused

landslides: a major disturbance to tropical forests. Science

205: 997-999.

Gamundí, I. 1971. Las Cyttariales sudamericanas. Darwiniana 16:

461-510.

Gentili, M. and P. Gentili. 1988. Lista comentada de los insectos

88
 asociados a las especies sudamericanas del género Nothofagus.

Monografías de la Academia Nacional de Ciencias Exactas, Físicas

y Naturales 4: 85-106.

Godoy, J.C. 1963. Fauna Silvestre. Evaluación de los Recursos

Naturales de la Argentina. Report, Consejo Federal de

Inversiones, Buenos Aires, Argentina.

Goodall, R.N.C. 1979. Tierra del Fuego. Ediciones Shanamaiim.

Buenos Aires, Argentina.

Grosse, J.A. 1979. Colonización en Aisén y Chiloé Continental.

Trapananda 3: 3-5.

Grosse, H. 1989. Renovales de Raulí, Roble, Coigüe y Tepa.

Expectativas de rendimiento. Ciencia e Investigación Forestal

3: 37-71.

Gutiérrez, E., J. Romaña, J. Fons, T. Sauras, and V.R. Vallejo.

In press. Ecología de Nothofagus betuloides y N. pumilio en

Tierra del Fuego. In J. Puigdefábregas, J. Frangi, F.G.

Novo, and A. Vila (eds.), The Natural Systems of Tierra

del Fuego and their Occupation by Man. Consejo Superior de

Investigación y Ciencia, Madrid.

Gutiérrez, E., V.R. Vallejo, J. Romañá, and Jaume Fons. 1991. The

subantarctic Nothofagus forests of Tierra del Fuego:

Distribution, Structure and Production. Oecologia Aquatica 10:

1-14.

Gutiérrez, J. 1989. Manejo de renovales nativos. Revista CORMA

207: 32-36.

Heusser, C.J. 1964. Some pollen profiles from the Laguna de San

89
 Rafael Area, Chile. In Ancient Pacific Floras, 10th Pacific

Science Congress Series. University of Hawaii Press, Honolulu,

pp. 95-115.

-----. 1987. Fire history of Fuego-Patagonia. Quaternary of

South America and Antarctic Peninsula 5: 93-109.

Heusser, C.J., J. Rabassa, A. Brandani, and R. Stuckenrath.

1988. Late-Holocene vegetation of the Andean Araucaria

Province of Neuquén, Argentina. Mountain Research and Development

8: 53- 63.

Holdgate, M.W. 1961. Vegetation and soils in the south Chilean

islands. J. Ecol. 49: 559-580.

Hosseus, C.K. 1915. Las cañas de bambú en las cordilleras del

sud. Boletín del Min. de Agr. Vol. 15. Buenos Aires.

Illies, H. 1970. Geología de los Alrededores de Valdivia y

Volcanismo y Tectónica en Márgenes del Pacífico en Chile

Meridional. Valdivia, Instituto de Geología y Geografía,

Universidad Austral de Chile.

Innes, J.L. 1992. Structure of evergreen temperate rain forest on

the Taitao Peninsula, southern Chile. J. Biogeog. 19: 555-562.

Kalela, E.K. 1941. Über die Holzarten und die durch die

klimatischen Verhältnisse verursachten Holzartenwechsel in den

Wäldern Ostpatagoniens. Ann. Acad. Sient. Fenn. Ser. A, 2: 5-

151.

Kamm, T. 1993. Sheep and trees are acting strangely at 'End of

the World.' Wall Street Journal 74(62): A1, A5

Kanamori, H. 1988. Importance of historical seismograms for

90
 geophysical research. In W.H.K. Lee, H. Meyers, and K.

Shimazaki (eds.), Historical Seismograms and Earthquakes of the

World, Academic Press, New York, pp. 16-33.

Kitzberger, T., and T.T. Veblen. Unpublished. Stand development

and recent vegetation trends along the northern Patagonian

forest-steppe gradient. Unpublished m.s.

Kitzberger, T., T.T. Veblen, and R. Villalba. Unpublished a.

Fire regimes along a rainforest to steppe gradient in northern

Patagonia: human and climatic influences. Unpublished m.s.

Kitzberger, T., T.T. Veblen, and R. Villalba. Unpublished b.

Tectonic influences on tree growth in northern Patagonia,

Argentina: The roles of climatic variation and substrate

stability. Unpublished m.s.

Kozlowski, T.T., P.J. Kramer, and S.G. Pallardy. 1991. The

Physiological Ecology of Woody Plants. Academic Press. San Diego.

Kuhnemann, O. 1976. Observaciones ecologicas sobre la vegetacion

marina y terrestre de la Isla de los Estados (Tierra del Fuego,

Argentina). ECOSUR, Argentina 3(6): 121-248.

LaMarche, V.C.Jr, R.L. Holmes, P.W. Dunwiddie, and L.G. Drew.

1979. Tree-ring chronologies of the southern hemisphere,

Chronology Series V. Tucson. Laboratory of Tree-Ring Research,

University of Arizona, Tucson, USA.

Lara, A. 1991. The dynamics and disturbance regimes of Fitzroya

cupressoides forests in the south-central Andes of Chile. Ph.D.

thesis, University of Colorado, Boulder, Colorado, U.S.A.

Lara, A. and T.T. Veblen. 1992. Forest plantations in Chile: A

91
 successful model? In A. Mather (ed.), Afforestation: Policies,

Planning and Progress. Belhaven Press, London, pp. 118-139.

Lawrence, D.B. and E.G. Lawrence. 1959. Glacier variations in

southern South America. Technical Report, Office of Naval

Research Contract 641-(04) to the American Geographical

Society.

Lechowicz, M.J. 1984. Why do temperate deciduous trees leaf out

at different times? Adaptation and ecology of forest

communities. Am. Nat. 124: 821-842.

Lizarralde, M., J. Escobar, and O. Biancciotto. 1990. El castor

(C. canadensis) en Tierra del Fuego: effectos de la alteración

ambiental y su aprovechamiento como recurso natural. Libro del

PEOAF, Museo del Fin del Mundo, Ushuaia, Argentina.

Lomnitz, C. 1970. Major earthquakes and tsunamis in Chile during

the period 1535 to 1955. Geologische Rundschau 59: 939-960.

Machoni, A. 1732. Las Siete Estrellas de la Mano de Jesús,

Tratado Histórico. Colegio de Asumpción, Córdoba, Spain.

Manabe, S. and R.T Wetherald. 1987. Large-scale changes of soil

wetness induced by an increase in atmospheric carbon dioxide.

J. Atmospheric Sci. 44: 1211-1235

Manion, P.D. 1981. Tree Disease Concepts. Prentice Hall,

Englewood Cliffs, New Jersey.

Markgraf, V. 1983. Late and postglacial vegetational and

paleoclimatic changes in subantarctic, temperate, and arid

environments in Argentina. Palynology 7: 43-70.

-----. 1987. Paleoclimates of the southern Argentine Andes.

92
 Current Research in the Pleistocene 4: 150-157.

Martín, C.D., M. Mermoz, and G. Gallopín. 1985. Impacto de la

ganadería en la cuenca del Rio Manso Superior. Report,

Administración de Parques Nacionales, Buenos Aires, Argentina.

Martinic, B.M. 1985. La ocupación y el impacto del hombre sobre

el territorio. In O. Boelcke, D.M. Moore, and F.A. Roig (eds.),

Transecta Botánica de la Patagonia Austral, Consejo Nacional

de Investigaciones Científicas y Técnicas, Buenos Aires, pp.

81-94 .

Mascareño, A. 1987. Evaluación de ensayos de semillación y

regeneración de Lenga (N. pumilio) bajo diferentes tratamientos

a la cama de semillas en la Reserva Forestal Trapananda,

Coyhaique. Thesis, Universidad Austral de Chile, Valdivia, Chile.

McQueen, D.R. 1976. The ecology of Nothofagus and associated

vegetation in South America. Tuatara 22: 38-68.

Miller, A. 1976. The climate of Chile. In W. Schwerdtfeger (ed.),

World Survey of Climatology. Vol. 12: Climates of Central and

South America. Elsevier, Amsterdam, pp. 113-131.

Moore, D.M. 1979. Southern oceanic wet-heathlands (including

magellanic moorland). In R.L. Specht (ed.), Heathlands and

Related Shrublands, Elsevier, Amsterdam, pp. 489-497

-----. 1983. Flora of Tierra del Fuego. Missouri Botanical Garden,

St Louis, Missouri.

Moreno, F.P. 1897. Reconocimiento de la región andina de la

República Argentina. Apuntes preliminares sobre una excursión a

los Territorios de Neuquén, Rio Negro, Chubut y Santa Cruz.

93
 Revista del Museo de La Plata 8: 1-180.

Mueller-Dombois, D. 1987. Natural dieback in forests. Bioscience

37: 575-583.

-----. 1988. Towards a unifying theory for stand-level dieback.

GeoJournal 17: 249-252.

Müller-Using, B. and F.M. Schlegel. 1981. The development of

seedlings of Chilean Nothofagus species in a shaded area. Plant

Research and Development 13: 152-184.

Musters, G.C. 1871. At Home with Patagonians: A Year's Wandering

over Untrodden Ground from the Straits of Magellan to the Rio

Nergo. John Murray, London, England.

Mutarelli, E.J. and E.N. Orfila. 1971. Observaciones sobre la

regeneración de lenga, Nothofagus pumilio (Poepp. et Endl.),

Oerst., en parcelas experimentales del lago Mascardi,

Argentina. Rev. Forestal Argentina 15: 109-115.

Mutarelli, E.J. and E.N. Orfila. 1973. Algunos resultados de las

investigaciones de manejo silivicultural que se realizan en los

bosques andino-patagónicos de Argentina. Rev. Forestal

Argentina 17: 69-74.

Oberdorfer, W. 1960. Pflanzensoziologische Studien in Chile Ein

Vergleich mit Europa. Flora et Vegetatio Mundi 2: 1-208.

Oliver, C.D. 1981. Forest development in North America

following major disturbances. For. Ecol. Manag. 3: 153-168.

Ordoñez, A. 1986. Germinación de las tres especies de Nothofagus

siempreverdes (Coigües) y variabilidad en la germinación de

procedencias de Coigüe común (N. dombeyi). Thesis, Universidad

94
 Austral de Chile, Valdivia, Chile.

Orfila, E. 1978. Misodendraceae de la Argentina y Chile. Buenos

Aires, Fundación Elias y Ethel Malamud. Serie Científica.

Ormazábal, C. 1992. Informe Nacional del PAF (Plan de Acción

Forestal). Unpublished report, Santiago.

Ortiz, J., R. Rodríguez, C. Marticorene, G. Riveros, H. Ibarra,

F. Peña, V. Jeréz, V.H. Ruiz, V. Quintana, J. Solervicens, J.

Artigas and E. Ugarte. 1993. Sitios prioritarios para la

conservación de la diversidad biológica en Chile. Documento

CONAF. Universidad de Concepción, Concepción, Chile.

Pearson, O.P. and A.K. Pearson. 1982. Ecology and biogeography of

the southern rainforests of Argentina. In M.A. Mares and H.H.

Genoways (eds.), Mammalian Biology in South America. Special

Publication Series, Pymatuning Laboratory of Ecology,

University of Pittsburg, Linesville Penssylvania 6: 129-142.

Peredo, H.L. 1987. Fitoparásitios en Nothofagus chilenos. Bosque

8: 105-107.

Perez, R. 1958. Recuerdos del Pasado: 1814-1860. Carlos de Vidts,

Santiago.

Petersen, J.G. 1988. Chilecomadia valdiviana (Philippi)

(Lepidoptera: Cossidae), asociado a Nothofagus pumilio (Poepp.

et Endl) Krasser (lenga) en la Región de Magallanes. Anales del

Instituto de la Patagonia 18: 51-55.

Peterson, C.J. and S.T.A. Pickett. 1991. Treefall and resprouting

following catastrophic windthrow in an old-growth hemlock-

hardwoods forest. Forest Ecology and Management 42: 205-217.

95
Pickett, S.T.A. and J.N. Thompson. 1978. Patch dynamics and the

size of nature reserves. Biol. Cons. 13: 27-37.

Pisano, V.E. 1973. Fitogeografía de la Península Brunswick,

Magallanes. I. Comunidades meso-higromórficas e higromórficas.

Anales del Instituto de la Patagonia 4: 141-206.

-----. 1978. Establecimientos de Nothofagus betuloides (Mirb.)

Blume (Coigue de Magallanes) en un valle en proceso de

deglaciación. Anales del Instituto de la Patagonia 9: 107-128.

Pisano, E. and M.J. Dimitri. 1973. Estudio ecológico de la región

continental sur del area andino-patagónica. Anales del

Instituto de la Patagonia 4: 207-271.

Povilitis, A. 1978. The Chilean Huemul Project (1975-76): Huemul

Ecology and Conservation. Ph.D. thesis, Colorado State

University, Fort Collins, Colorado, U.S.A.

Premoli, A.C. 1991. Morfología y capacidad germinativa en

poblaciones de Nothofagus antarctica (Foster)Oerst. del

noroeste andino patagónico. Bosque 12: 53-59.

-----. Umpublished. Genetic variation in two widespread and a

geographically restricted species of South American Nothofagus.

Unpublished m.s.

Puigdefábregas, J., O. Bianciotto, G. del Barrio, G. Gallart, N.

Clotet, M. Alogia and L. Ferres. In press. Respuesta del bosque

subantánartico frente a perturbaciones naturales. In

Puigdefábregas, J., J. Frangi, F.G. Novo, and A. Vila (eds.),

The Natural Systems of Tierra del Fuego and their Occupation by

Man. Consejo Superior de Investigación y Ciencia, Madrid.

96
Raedeke, K. 1979. Population dynamics and socioecology of the

guanaco (Lama guanicoe) of Magallanes, Chile. Ph.D. thesis,

University of Washington, Seattle, Washington, U.S.A.

Ramilo, E. 1985. Situación y manejo del ciervo colorado en los

Parques Nacionales Lanín y Nahuel Huapi. Report, Administración

de Parques Nacionales, Buenos Aires, Argentina.

Ramírez, C., M. Correa, H. Figueroa, and J. San Martín. 1985.

Variación del hábito y habitat de Nothofagus antarctica en el

centro sur de Chile. Bosque 6: 55-73.

Ramírez, C. and H. Figueroa. 1985. Fitosociología de los

Nothofagus de la zona higromórfica chilena. Bosque 6: 127-132.

Ramírez, C., R. Godoy, W. Eldridge, and N. Pacheco. 1981. Impacto

ecológico del ciervo rojo sobre el bosque de Olivillo en Osorno,

Chile. Anales del Museo de Historia Natural 14: 197- 212.

Read, J. and R.S. Hill. 1985. Dynamics of Nothofagus-dominated

rainforest on mainland Australia and lowland Tasmania.

Vegetatio 63: 67-78.

Rebertus, A.J. and T.T. Veblen. 1993a. Partial wave formation in

old-growth Nothofagus forests of Tierra del Fuego. Bull. Torr.

Bot. Cl. 120: 461-470.

Rebertus, A.J. and T.T. Veblen. 1993b. Structure and tree-fall

gap dynamics of old-growth Nothofagus forests in Tierra del

Fuego, Argentina. J. Veg. Sci. 4: 641-654.

Rebertus, A.J., T.T. Veblen, and T. Kitzberger. 1993. Gap

formation and dieback in Fuego-Patagonian Nothofagus forests.

Phytocoenologia 23: 581-599.

97
Richter, L.L. and J.L. Frangi. 1992. Bases ecológicas para el

manejo del bosque de Nothofagus pumilio de Tierra del Fuego.

Revista de la Facultad de Agronomía 68: 35-52.

Rocuant, L. 1984. Efecto de giberelina y de tiourea en la

germinación de semillas: especies del género Nothofagus. Bosque

5: 53-58.

Roig, F.A. J. Anchorena, O. Dollenz, A.M. Faggi, and E. Méndez.

1985. Las comunidades vegetales de la transecta botánica de la

Patagonia austral. In O. Boelcke, D.M. Moore, and F.A. Roig,

(eds.), Transecta Botánica de la Patagonia Austral, Consejo

Nacional de Investigaciones Científicas y Técnicas, Buenos

Aires, pp. 350-519.

Rothkugel, M. 1916. Los Bosques Patagónicos. Ministerio de

Agricultura, Buenos Aires, Argentina.

Runkle, J.R. 1985. Disturbance regimes in temperate forests. In

S.T.A. Pickett and P.S. White (eds.), The Ecology of Natural

Disturbance and Patch Dynamics, Academic Press, New York, pp.

17-33

Runkle, J.R. 1990. Gap dynamics in an Ohio Acer-Fagus forest and

speculations on the geography of disturbance. Can. J. For. Res.

20: 632-641.

Rusch, V. 1987. Estudio sobre la regeneración de la lenga en la

Cuenca dle Río Manso Superior, Río Negro. Unpubl. report,

Consejo Nacional de Ciencia y Tecnología, Buenos Aires.

-----. 1992. Principales limitantes para la regeneración de la

lenga en la zona N.E. de su área de distribución. Variables

98
 ambientales en claros del bosque. In J. Bava and J.

Schmalz (eds.), Actas Seminario de Manejo Forestal de la

Lenga y Aspectos Ecológicos Relacionados. CIEFAP,

Publicación Técnica 8. Esquel, pp. 61-73.

-----. 1993. Altitudinal variation in the phenology of Nothofagus

pumilio in Argentina. Rev. Chil. His. Nat. 66: 131-141.

Sakai, A., D.M. Paton, and P. Wardle. 1981. Freezing resistance

of trees of the South temperate zone, especially subalpine

species of Australasia. Ecol. 62: 563-570.

Sakai, A. and P. Wardle. 1978. Freezing resistance of New Zealand

trees and shrubs. N.Z. J. Ecol. 1: 51-61.

Schlegel, F.S., T.T. Veblen, and B. Escobar R. 1979. Estudio

ecológico de la estructura, composición, semillación y

regeneración del bosque de Lenga (Nothofagus pumilio) XI Región.

Universidad Austral de Chile, Facultad de Ingeniería Forestal,

Serie Técnica.

Schmidt, H. and A. Urzúa. 1982. Transformación y manejo de los

bosques de lenga en Magallanes. Ciencias Agrícolas (Universidad

de Chile, Santiago) No. 11, pp. 1-62.

Simonetti, J.A. and J.J. Armesto. 1991. Conservation of temperate

ecosystems in Chile: Coarse versus fine-filter approaches. Rev.

Chil. His. Nat. 64: 615-626.

Singer, R. 1971. Forest mycology and forest communities in South

America. II. Mycorrhiza sociology and fungus succession in the

Nothofagus dombeyi-Austrocedrus chilensis woods of Patagonia.

Proc. of the North American Conference on Mycorrhizae. USDA

99
 Misc. Publ. 1189.

Singer, R. and J.H. Morello 1960. Ectotrophic forest tree

mycorrhiza and forest communities. Ecol. 41: 549-551.

Singer, R. and M. Moser. 1965. Forest mycology and forest

communities in South America. Mycopathologia et Mycologia

Applicata 26: 9-189.

Smulders, A.M. 1988. Estudio del ritmo de crecimiento de N.

alpina, N. dombeyi y Persea lingue durante sus primeros años de

vida. Thesis, Universidad Austral de Chile, Valdivia, Chile.

Stewart G.H., Rose, A.B., and T.T. Veblen. 1991. Forest

development in tree-fall gaps in old-growth beech (Nothofagus)

forests, New Zealand. Journal of Vegetation Science. J. Veg.

Sci. 2: 679-690.

Tortorelli, L.A. 1947. Los Incendios de Bosques en la Argentina.

Ministerio de Agricultura, Buenos Aires, Argentina.

-----. 1956. Maderas y bosques argentinos. Editorial ACME, Buenos

Aires.

Tukhanen, S. 1992. The climate of Tierra del Fuego from a

vegetation geographical point of view and its ecoclimatic

counterparts elsewhere. Acta Bot. Fenn. 145: 1-64.

Veblen, T.T. 1979. Structure and dynamics of Nothofagus forests

near timberline in south-central Chile. Ecol. 60: 937-945.

-----. 1982a. Growth patterns of Chusquea bamboos in the

understory of Chilean Nothofagus forests and their influences

in forest dynamics. Bull. Torrey Botanical Club 109: 474-487.

-----. 1982b. Regeneration patterns in Araucaria araucana forests

100
 in Chile. J. Biogeog. 8: 211-247.

-----. 1985a. Forest development in tree-fall gaps in the

temperate rain forests of Chile. National Geographic Research

1: 161-184.

-----. 1985b. Stand dynamics in Chilean Nothofagus forests. In

S.T.A. Pickett and P.S. White (eds.), The Ecology of Natural

Disturbance and Patch Dynamics, Academic Press, New York, pp.

35-51.

-----. 1989a. Nothofagus regeneration in treefall gaps in

northern Patagonia. Can. J. For. Res. 19: 365-371.

-----. 1989b. Tree regeneration responses to gaps along a

transandean gradient. Ecol. 70: 543-545.

-----. 1992. Regeneration dynamics. In D.C. Glenn-Lewin, R.K.

Peet, and T.T. Veblen (eds.), Plant Succession: Theory and

Prediction. Chapman and Hall, London. Pp. 152-187.

Veblen, T.T. and P.B. Alaback. In press. A comparative review of

forest dynamics and disturbance in the temperate rain forests

of North and South America. In: R.G. Lawford, P. Alaback and

E.R. Fuentes (eds.), High Latitude Rain Forests and Associated

Ecosystems of the West Coast of the Americas: Climate,

Hydrology, Ecology and Conservation. Springer-Verlag, New York.

Veblen, T.T. and D.H. Ashton. 1978. Catastrophic influences on

the vegetation of the Valdivian Andes. Vegetatio 36: 149-167.

Veblen, T.T., D.H. Ashton, S. Rubulis, D.C. Lorenz, and M.

Cortés. 1989a. Nothofagus stand development on in-transit

moraines, Casa Pangue Glacier, Chile. Arct. Alp. Res. 21: 144-

101
 155.

Veblen, T.T., D.H. Ashton, F.M. Schlegel, and A.T. Veblen. 1977.

Plant succession in a timberline depressed by volcanism in

south-central Chile. J. Biogeog. 4: 275-294.

Veblen, T.T., C. Donoso Z, F.M. Schlegel, and B. Escobar. 1981.

Forest dynamics in south-central Chile. J. Biogeog. 8: 211-247.

Veblen, T.T., T. Kitzberger, and A. Lara. 1992a. Disturbance and

forest dynamics along a transect from Andean rain forest to

Patagonian shrublands. J. Veg. Sci. 3: 507-520.

Veblen, T.T. and D. C. Lorenz. 1987. Post-fire stand development

of Austrocedrus-Nothofagus forests in Patagonia. Vegetatio 73:

113-126.

Veblen, T.T. and D.C. Lorenz. 1988. Recent vegetation changes

along the forest/steppe ecotone in northern Patagonia. Annals

of the Association of American Geographers 78: 93-111.

Veblen, T.T. and V. Markgraf. 1988. Steppe expansion in

Patagonia? Quat. Res. 30: 331-38.

Veblen, T.T., Mermoz, M. Martín, C., and E. Ramilo. 1989b.

Effects of exotic deer on forest regeneration and composition

in northern Patagonia. J. Appl. Ecol. 26: 711-724.

Veblen, T.T., M. Mermoz, C. Martín, and T. Kitzberger. 1992b.

Ecological impacts of introduced animals in Nahuel Huapi

National Park, Argentina. Cons. Biol. 6: 71-83.

Veblen, T.T., F.M. Schlegel, and B. Escobar. 1980. Structure

and dynamics of old-growth Nothofagus forests in the Valdivian

Andes, Chile. J. Ecol. 68: 1-31.

102
Veblen, T.T., F.M. Schlegel, and J.V. Oltremari. 1983. Temperate

broad-leaved evergreen forests of South America. In J.D.

Ovington (ed.), Temperate Broad-Leaved Forests, Elsevier,

Amsterdam, pp. 5-31 .

Vera, O.J. 1985. Evaluación de intervenciones silivícolas en un

renoval mixto de Lenga (N. pumilio) y Coigüe (N. dombeyi)

ubicado en la Reserva Forestal Coyhaique. Thesis, Universidad

Austral de Chile, Valdivia, Chile.

Waters, J.W., L. Froidevaux, W.G. Read, G.L. Manney, L.S. Elson,

D.A. Flower, R.F. Jarnot, and R.S. Harwood. 1993. Stratospheric

ClO and ozone from the Microwave Limb Sounder on the Upper

Atmosphere Research Satellite. Nature 362: 597-602.

Welch, C. 1988. Phytophagous insects on deciduous Nothofagus in

Chile and Argentina. Monografías de la Academia Nacional de

Ciencias Exactas, Físicas y Naturales 4: 107-114.

Werner, J. 1987. Determinación de períodos óptimos de

estratificación para semillas de diferentes procedencias de

Raulí. Thesis, Universidad Austral de Chile, Valdivia, Chile.

Willis, B. 1914. El Norte de la Patagonia. Dirección de Parques

Nacionales, Buenos Aires, Argentina.

Wright, J.E. 1987. Interrelaciones entre Macromycetes (Fungi) y

Nothofagus. Monografías de la Academia Nacional de Ciencias

Exactas, Físicas y Naturales 4: 135-152.

Young, S.B. 1972. Subantarctic rain forest of Magellanic Chile:

distribution, composition, and age growth rate studies of

common forest trees. Antarctic Reserch Series 20: 307-322.

103
Yudelevich, M., C. Brown, H. Elgueta, and S. Calderon. 1967.

Clasificación Preliminar del Bosque Nativo de Chile. Informe

Técnico No. 27. Instituto Forestal de Chile. Santiago.

Table 10.1 Common tree species of the major forest zones of

southern Chile and southwestern Argentina.

Valdivian Patagonian Magellanic

Cool Temperate

Rainforest Rainforest Rainforest

Nothofagus Forests

and woodlands

Nothofagus alpina* x

Nothofagus obliqua* x

Laurelia sempervirens x

Amomyrtus meli x

Crinodendron hookerianum x

Eucryphia cordifolia x

Persea lingue x

Aextoxicon punctatum x

Gevuina avellana x

Fitzroya cupressoides# x

Luma apiculata x

Podocarpus saligna# x

104
Nothofagus nitida x x

Laurelia philippiana x x

Amomyrtus luma x x

Pseudopanax laetevirens x x

Chusquea bamboos x x

Caldcluvia paniculata x x

Saxegothaea conspicua# x x

Weinmannia trichosperma x x

Nothofagus dombeyi x x

Podocarpus nubigena# x x

Maytenus magellanica x x x

Drimys winteri x x x

Pilgerodendron uviferum# x x x

Embothrium coccineum x x x

Nothofagus betuloides x x x

Nothofagus antarctica* x x x

Nothofagus pumilio* x x x

Araucaria araucana#

Austrocedrus chilensis#

105
Lomatia hirsuta

Deciduous broadleaved species are indicated by * and evergreen

conifers by #. All others are evergreen broadleaved trees.

106
Table 10.2 Phenological behavior of Nothofagus species in southern Chile. Data
from: Donoso and Cabello 1978, Becker 1981, and Mascareño 1987.

Caracter Spp. J F M A M J J A S O N D
Bud burst N. alpina $$$$$
N. obliqua $$$$$
N. pumilio $$$$$
N. antarctica
N. dombeyi $$$$$$$$$
N. betuloides
N. nitida $$$$$$$$$
J F M A M J J A S O N D
Leaf fall N. alpina $$$$$
N. obliqua $$$$
N. pumilio $$$$$
N. antarctica
N. dombeyi
N. betuloides
N. nitida
J F M A M J J A S O N D
Flowering N. alpina $$$$$$$$$
N. obliqua $$$$$
N. pumilio
N. antarctica
N. dombeyi $$$$$
N. betuloides
N. nitida $$$$$$$$$
J F M A M J J A S O N D
Seed fall N. alpina $$$$$
N. obliqua $$$$$$$
N. pumilio $$$$$
N. antarctica $$$$$
N. dombeyi $$$$$$$$$$$$$$$$$
N. betuloides $$$$$
N. nitida $$$$$$$$$

107
Table 10.3 Latitudinal variation in germination capacity of Nothofagus

obliqua, N. alpina and N. dombeyi for control and optimum stratification

treatments. Data from: Ordoñez 1986, Donoso 1987, and Werner 1987.

Species and origin Latitude S Germination capacity

(percent)

S)))))))))))))))))))))))))))))))Q

Control Optimum stratification

treatment (days)
S))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))Q

Nothofagus obliqua

Coastal Range 33° 07' 18.7 52.0 (60)

Coastal Range 34° 12' 26.7 59.3 (60)

Andean Cordillera 35° 15' 32.7 53.3 (90)

Andean Cordillera 36° 20' 33.3 42.0 (90)

Andean Cordillera 36° 30' 37.3 52.7 (90)

Andean Cordillera 38° 20' 30.7 64.7 (90)

Andean Cordillera 39° 20' 5.0 17.0 (60)

Andean Cordillera 40° 20' 16.7 20.0 (30)

Andean Cordillera 41° 08' 23.3 14.7 (60)

Nothofagus alpina

Andean Cordillera 35° 40' 45.3 80.0 (60)

Andean Cordillera 36° 45' 42.6 62.0 (60)

Andean Cordillera 38° 00' 30.6 38.0 (60)

108
 Andean Cordillera 39° 20' 7.3 10.6 (60)

Andean Cordillera 39° 50' 23.3 24.0 (60)

Andean Cordillera 40° 25' 51.3 50.6 (60)

Coastal Range 40° 25' 54.0 43.3 (60)

Nothofagus dombeyi

Andean Cordillera 36° 52' 51.2 61.7 (90)

Andean Cordillera 37° 35' 19.0 71.0 (90)

Coastal Range 39° 08' 2.5 17.0 (90)

Andean Cordillera 39° 36' 5.2 22.2 (90)

Coastal Range 40° 16' 0.5 10.0 (90)

109
Table 10.5 Gap and treefall characteristics in Nothofagus-dominated forests in South America and New Zealand (N.Z.). TDF stands for Tierra del Fuego. Locations: Turnover

times or rotation periods are based on canopy gap areas. No distinction was made between N. pumilio and N. betuloides treefalls in the Tierra del Fuego study. Separate treefall modes

are given for N. fusca and N. menziesii.

Uproot: Median Gap Size (m2) Turnover Dominant

Species Location Windsnap Canopy Expanded Time (yrs) filler Source

N. pumilio Tierra del Fuego 1.6 104 268 346-519 self-maintaining Rebertus and Veblen 1993b

N. pumilio-N. Tierra del Fuego 1.3 61 153 331-415 both species coexist Rebertus and Veblen 1993b

betuloides

N. betuloides Beagle Channel, TDF 0.9 self-maintaining Rebertus and Veblen 1993 b

N. betuloides Beagle Channel, TDF 1.7 106 208 320-448 Drimys winteri Rebertus and Veblen 1993 b

(with Drimys)

110
N. dombeyi Chilean Lake District 0.5 --- 542 392 Chusquea and Laurelia philippiana Veblen 1985a

(with others)

N. dombeyi Argentina Lake District 2.4 --- 403-603 --- N. dombeyi and Austrocedrus coexist Veblen 1989a

(with Austrocedrus)

N. antarctica Argentina Lake District 4.1 --- --- --- N. dombeyi Veblen 1989a

(with N. dombeyi)

N. fusca-N. Station Crk., N.Z. 0.5, 1.4 68 260 both species coexist Stewart et al. 1991

menziesii Fergies Bush, N.Z. 0.5, 0.7 140 295

Rough Crk., N.Z. 0.4, 0.5 107 353

111
Table 10.6 Standing crops in Nothofagus forests on Tierra del Fuego. All stands are
considered old-growth except the N. pumilio stand on an unstable slope at 555 m

elevation in which the oldest trees were <100 years old.

Forest type Latitude Elevation Standing crop (mt/ha)

(S) (m) Aboveground Belowground Total

S))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))Q

N. antarctica1 54° 100 217.1 30.5 247.6

N. pumilio1 55° 100 431.9 66.3 498.2

N. pumilio2 55° 260 283.9 n.d. n.d.

N. pumilio2 55° 375 265.1 n.d. n.d.

N. pumilio2 55° 555 126.9 n.d. n.d.

N. betuloides1 55° 100 700.9 166.9 867.8

N. betuloides2 55° 260 468.0 n.d. n.d.

2
N. betuloides 55° 375 246.6 n.d. n.d.
2
N. betuloides 55° 555 311.6 n.d. n.d.

1 2
Sources: Frangi and Richter (in press); Gutiérrez et al. (in press)

112
Table 10.7 Annual fine-litter production in Nothofagus forests in southern Chile and
Argentina. Fine litter includes leaves, twigs < 1 cm diameter, and reproductive

structures.

Vegetation type Latitude Altitude Total Chusquea

(m) (mt/ha) (% of total)

S))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))))

Old-growth Nothofagus 40o 810 5.61 5.5

dombeyi- Saxegothaea
conspicua-Laurelia

philippiana
Old-growth N. dombeyi- 40o 1100 5.42 n.a.

N. alpina-S. conspicua-
L. philippiana

Bamboo thicket of 40o 700 3.91 100.0

Chusquea culeou
Old-growth N. dombeyi- 40o 1060 5.01 1.3

N. alpina-N. pumilio
Second-growth N. alpina 40o 700 5.41 12.3

Second-growth N. dombeyi- 40o 60 7.11 6.4

N. obliqua
Second-growth N. obliqua 40o 50 7.41 9.7
Laurelia sempervirens
Old-growth N. antarctica 54o 100 3.03 n.a.

Old-growth N. pumilio 55o 100 3.73 n.a.

Old-growth N. betuloides 55o 100 2.73 n.a.
Old-growth N. pumilio 55o 260 2.04 n.a.
Old-growth N. pumilio 55o 375 2.44 n.a.
Second-growth N. pumilio 55o 555 2.14 n.a.

113
Old-growth N. betuloides 55o 25 2.24 n.a.
o 4
Old-growth N. betuloides 55 150 2.4 n.a.
o 4
Old-growth N. betuloides 55 335 3.4 n.a.

Sources: 1Veblen 1982a; 2Burschel et al. 1976; 3Frangi and Richter in press; and 4Gutiérrez
et al. in press.

114
Table 10.4 Insects found on Nothofagus spp. in southern Chile and Argentina. Numbers indicate the number of species per family (after Gentili and

Gentili 1988). Species reported as pests for the different insect families are listed after Carrillo and Cerda (1987).

N. antarctica N. pumilio N. obliqua N. alpina N. nitida N. dombeyi/ N. betuloides Pest species

FOLIAGE CONSUMERS
Coleoptera
Scarabeidae 1 4 2 Hylamorpha elegans

Sericoides germaini
Curculionidae 3 1 3 5

Crysomelidae 1

Lepidoptera

Hemileucidae Automeris spp.

Ormiscodes cinnamomea
Polythysana rubrescens

Lasiocampidae 1 Macromphalia spp.

Geometridae (various insect spp. not determined) Omaguacua longibursae

Hymenoptera

Pergidae 1 1 Cerospastus volupis

LIVE AND DEAD WOOD BORERS, BARK AND CAMBIUM AND ROOT FEEDERS

Coleoptera
Anobiidae 1 Callymmaderus capucinus Bostrichidae
Dexicrates robustus

Buprestidae 3 1 2 Epistomentis picta

Pterobothris corrosus
Cerambycidae 10 7 6 2 20 Acanthidonera cumminigi

Calydon submetallicum

115
 Callideriphus laetus

Callisphyris spp.

Cheloderus peñai

Holopterus chilensis
Lautarus concinnus

Microplophorus magellanicus
Oxipeltus quadrispinosus
Cucujidae 1 1

Curculionidae 2 2 5 Aegorhinus vitulus

Empleurodes dentipes

Rhyephenes spp.

Dermestidae 1

Elateridae 1 2
Limexylonidae 1 1
Lucanidae 1 1 3

Lyctidae Lyctus brunneus

Scrabeidae 1 2 2 Hyalomorpha elegans

Scolytidae 1 Sinophloeus porteri

Isoptera
Kalotermitidae Cryptotermes brevis

Lepidoptera
Cossidae 2 1 Chilecomadia moorei
Chilecomadia valdiviana

Hepialidae 1
Noctuidae various genera
Opostegidae 1 Opostega spp.

116
FRUIT AND SEED FEEDERS

Lepidoptera

Heterobathmiidae 1 1 1

Oecophoridae Perzelia spp.

N. antarctica N. pumilio N. obliqua N. alpina N. nitida N. dombeyi/ N. betuloides Pest species

SAPSUCKERS

Heteroptera
Coreidae 1

Homoptera
Aachilidae 1

Aphididae 1 8 4
Cicadidae 1 1
Cixidae 1 1

Conchaspididae 1
Delphacidae (host species not specified)

Diaspididae 1 1
Ericoccidae 2 4 2 6

Margarodidae 1 1
Membracidae 1
Phyloxeridae 1

Thysanoptera
Aeolothripidae 1 1

Thripidae 1

GALL FORMERS

Coleoptera

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Curculionidae 1 1 1

Hymenoptera

Figitidae 1
Pteromalidae 1 1 1

LEAF MINERS
Lepidoptera

Heterobathmidae 1 1 3
Tortricidae 1

OTHER UNSPECIFIED RELATIONSHIPS

Coleoptera

Buprestidae 1 1 2
Cerambicidae 1 1 1 5
Curculionidae 1 2 2

Elateridae 4 2

FIGURE CAPTIONS

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FIGURE 10.1. Location map and the latitudinal distribution of Nothofagus in southern Chile and

Argentina. After: Donoso (1987).

FIGURE 10.2. Mean monthly temperature and precipitation for three stations within the ranges of

southern Chilean and Argentinean Nothofagus forests. Concepción and Punta Arenas are located on

the coast of Chile and Bariloche is on the eastern side of the Andes in Argentina. Note the

relatively high degree of seasonal temperature variation at Bariloche given its latitude.

FIGURE 10.3. Major vegetation zones of southern South America.

FIGURE 10.4. Leaves of the Nothofagus spp. occurring in southern Chile and Argentina. After

Donoso (1974)

FIGURE 10.5. Photograph of Nothofagus antarctica showing its typically shrubby form near the

ecotone with the Patagonian steppe at c. 40o S in Argentina. These individuals are

sprouting from the base seven years after being burned.

FIGURE 10.6. Photograph of Nothofagus obliqua in a c. 80-year old second-growth stand at c. 40oS

in Argentina. This stand has been heavily used by livestock.

FIGURE 10.7. Photograph of Nothofagus dombeyi showing its typical buttressed form. Trunk diameter

at breast height of this individual is 2.5 m.

FIGURE 10.8. Photograph of a pure N. dombeyi stand at c. 40oS in Argentina showing the typically

dense and uniform understory of 3 to 5-m tall Chusquea culeou bamboo beneath the relatively

open tree canopy.

FIGURE 10.9. Photograph of Nothofagus nitida with Gunnera chilensis at c. 47oS on glacial outwash near

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the Pacific coast of Chile.

FIGURE 10.10. Photograph of Nothofagus pumilio showing gap-phase regeneration (foreground)

following a treefall on Tierra del Fuego, Argentina.

FIGURE 10.11. Time series of photographs of a landslide on Lake Todos Los Santos, Chile triggered

by the 1960 Valdivian earthquake: (a) bare site six years following the landslide; (b) the same

site dominated by Gunnera chilensis in 1976; and (c) the same site dominated by Nothofagus dombeyi

in 1994.

FIGURE 10.12. Photograph of a stand of Nothofagus dombeyi and Austrocedrus chilensis at c. 40oS

in Argentina where the 1960 Valdivian earthquake caused extensive tree mortality.
FIGURE 10.13. Tree-ring width chronologies of Austrocedrus chilensis from two stands at c. 40oS in

Argentina where past earthquakes have affected the growth patterns of the surviving trees. The

numbers of tree cores included in the chronologies at various years are given in parentheses. In

both stands most of the trees killed in 1960 were Nothofagus dombeyi. Site F-2 is depicted in

Fig. 10.12. From: Kitzberger, Veblen, and Villalba unpublished m.s.

FIGURE 10.14. Photograph of a snow avalanche path through Nothofagus pumilio forest on Tierra del

Fuego.

FIGURE 10.15. Photograph of an even-aged Nothofagus dombeyi stand which originated after a

fire c. 80 years ago at c. 40oS in Argentina.

FIGURE 10.16. Photograph of dieback of Nothofagus pumilio near the steppe ecotone at c. 40oS in

Argentina.

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 FIGURE 10.17. Time series of photographs showing a treefall gap in a Nothofagus dombeyi-Laurelia
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philippiana-Saxegothaea conspicua forest in Chile being filled in by Chusquea culeou bamboo and

Laurelia philippiana: (a) 1977, one year after the treefall; (b) 1979; and (c) 1983. Note that in

each photograph a person is touching the same wind-snapped stump (indicated by the arrow).

FIGURE 10.18. Photograph of an unfilled treefall gap in a Nothofagus dombeyi forest in Argentina

where browsing by exotic deer have prevented tree regeneration.

FIGURE 10.19. Photograph of Nothofagus pumilio forests which have been drowned (center foreground)

by the damming activities of introduced beavers on Tierra del Fuego.

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