The Conifers of Chile: an Overview of their Distribution and Ecology

Martin F. Gardner Antonio Lara

Royal Botanic Garden Edinburgh Facultad de Ciencias Forestales
20A Inverleith Row Universidad Austral de Chile
Edinburgh EH3 5LR, UK Casilla 567
Valdivia, Chile

Keywords: Araucariaceae, Cupressaceae, Podocarpaceae, mesic, regeneration,

sclerophyllous vegetation, Valdivian rainforests

Abstract
The nine indigenous conifers of Chile are Araucaria araucana
(Araucariaceae), Austrocedrus chilensis, Fitzroya cupressoides, Pilgerodendron
uviferum (Cupressaceae), and Lepidothamnus fonkii, Podocarpus nubigenus, P.
salignus, Prumnopitys andina and Saxegothaea conspicua (Podocarpaceae). In Chile
these conifers have a 2,100 km north-south range where the annual precipitation can
vary in the north Mediterranean region from less than 1,000 mm to in excess of
4,000 mm in the southern magellanic rainforests. The Chilean conifers include some
of the tallest and oldest forest trees; Fitzroya is one of the oldest living plants in the
world, estimated to live in excess of 3,600 years. Many of these conifers have a
combination of slow growth rates, infrequent seed production and highly prized
wood and have therefore found it difficult to survive the non-sustainable exploits of
mankind.

INTRODUCTION
The 32 conifer species found in South America are represented by the following
nine genera: Araucaria (Araucariaceae), Austrocedrus, Fitzroya, Pilgerodendron
(Cupressaceae), Lepidothamnus, Podocarpus, Prumnopitys, Retrophyllum, and
Saxegothaea (Podocarpaceae). All of these except for Retrophyllum occur in Chile. It is
perhaps surprising that of the world’s 630 conifer species (Farjon, 2001), a continent the
size of South America, with such a diversity of mesic forests, contains less than 20% of
the world’s conifer species. Save for Araucaria angustifolia, little is known about the
ecology of the tropical conifer species of South America (Veblen et al., 1995); there is
also a great paucity in knowledge concerning their distributions.
In contrast, much more is known about the conifers in temperate South America,
especially those native to Chile. Of the nine indigenous Chilean conifers, eight also occur
across the Andes into Argentina (the exception being Podocarpus salignus), in most cases
the main distribution being in Chile. The conifer species in Chile have a latitudinal range
from 32°29′ to 55°30′ S and from sea-level to the Andean tree-line (Veblen, 1995).
Within this 2,100 km north-south latitudinal range, the climate varies from an annual
precipitation in excess of 3,000 mm in the Magellanic moorland and rainforests and the
Valdivian rainforests, to less than 2,000 mm in central Chile’s Mediterranean climate
region. Within a relatively small area of the Valdivian rainforests in southern Chile, all
nine conifer species occur, occupying a range of contrasting habitats from the coastal
Patagonian bogs to the well-drained volcanic ash slopes of the Andes.

THE CUPRESSACEAE
With a distribution throughout both hemispheres, the Cupressaceae is the most
widespread family of extant southern hemisphere conifers (Hill, 1995). In South America
the family is only found in Chile and Argentina where it is represented by the monotypic
genera Austrocedrus, Fitzroya and Pilgerodendron. The latter genus has recently been
considered congeneric with Libocedrus of New Caledonia and New Zealand on the basis
of molecular evidence (Gadek et al., 2000; Quinn and Price, this volume), but we retain
the name Pilgerodendron here.

Proc 4th IS on Conifer

Ed: R.R. Mill
Acta Hort 615, ISHS 2003 165
 In Chile, Austrocedrus chilensis spans c.1,100 km, from latitude 32°29′ to 43°38′
S (Dodd and Rafll, 1995) and of all the Chilean conifer species it has the most northerly
distribution, at San Felipe in the Region of Valparaíso. Within its native range it occurs
on steep-sided slopes from 300-1,850 m altitude (Dodd and Rafll, 1995) in scattered
populations but forms more sporadic populations in the north of its range. In common
with some other genera of the Cupressaceae (Cupressus and Tetraclinis in the northern
hemisphere and Callitris and Actinostrobus in the southern hemisphere), Austrocedrus is
capable of withstanding xeric conditions. In the Andes mountains, where Mediterranean
Chilean conifers occur, precipitation varies from less than 1,000 mm in the north to in
excess of 2,000 mm in the south of that region. In northern areas and in some Andean
locations Austrocedrus is associated with sclerophyllous species (Schmithüsen, 1960;
Donoso, 1982, 1996). An example of an Andean location is Parque Nacional Laguna de
Laja (37°24′S). Here at an altitude of 1,150 m where Austrocedrus is abundant, co-
dominant sclerophyllous vegetation includes Lomatia hirsuta, Quillaja saponaria,
Maytenus boaria, Orites myrtoidea, Fabiana imbricata, and Azara petiolaris. At higher
altitudes in this same location, Austrocedrus forms extensive pure stands. In wetter or
polar aspects, it is commonly associated with Nothofagus obliqua. At middle altitudes in
the Mediterranean Andes its association with N. obliqua and N. glauca is frequent
(Donoso, 1981, 1982, 1993). It is worth noting that A. chilensis forests are more abundant
and continuous between 36°30′ and 42°35′ on the east side of the Andes in Argentina
where precipitation is low (Havrylenko, 1989; Donoso, 1983). Older trees can range from
500 to 1,000 years of age (LaMarch et al., 1979). Recent dendrochronological research on
trees in the north of its range at San Gabriel (33°47′S), which barely reach a height of
three metres, have recorded the oldest to be 850 years (Aravena, pers. commun.).
Austrocedrus has a very limited range in the Pacific coastal mountains where it
occurs from the Cordillera de Nahuelbuta from 37°47′ to c. 40°30′S (Veblen and
Schlegel, 1982). At El Manzano (37°47′S) and Cerro Adencul (38°14′S), Austrocedrus is
confined to the margins of rivers where it is associated with Podocarpus salignus.
Many populations of Austrocedrus have at one time or another been subject to fire
damage (both man-made and natural); its ability to regenerate post-fire damage is good
although grazing by deer and cattle has had a devastating effect. Seeds are often severely
damaged by lepidopteran larvae, sometimes affecting the majority of the population.
Some populations have suffered from root rot caused by a fungal pathogen; this
condition, known as “mal del ciprés”, was first identified in the mid-1940s and typically
causes dead patches in populations of up to two hectares (Varsavsky et al., 1975).
In contrast to Austrocedrus chilensis, Fitzroya cupressoides and Pilgerodendron
uviferum (= Libocedrus uvifera) are both mesic species, occurring in some of the wettest
climates in Chile – from 2,000 to 6,000 mm of rain per annum. Pilgerodendron is a slow-
growing, dioecious conifer, capable of attaining a height of 40 m. It occurs between sea-
level and 1,000 m in a discontinuous north-south distribution of 1,500 km, from 39°35′S),
in the north (Martínez, 1981) to Tierra del Fuego (55°30′S), making it the most southerly
occurring conifer in the world. Typically, it is associated with Fitzroya in the coastal
cordilleras on exposed summits of the Cordillera Pelada and Cordillera de Piuché
(Chiloé) on poorly drained, thin gley soils (Peralta, 1978; Donoso, 1993). Other
commonly associated species include Nothofagus nitida, Tepualia stipularis and Drimys
winteri. Pilgerodendron is a frequent component of the southern Andes and the coastal
archipelago. There are large stands in the Andes of Palena, Aysén and Magellanes where
a type of wetland known as millines prevails (Innes, 1991; Donoso, 1993). Two woody
co-dominant species include Nothofagus betuloides and Drimys winteri with a ground
flora which includes Acaena pumila, Astelia pumila, Azorella caespitosa,
Marsippospermum grandiflorum, Myrteola nummularia and Perezia magellanica
(Moore, 1968).
In the low-lying areas of northern Chiloé and on the adjacent mainland (at
altitudes of less than 10 m), Pilgerodendron is locally abundant in sphagnum bogs, many
of which have been subject to fire damage (some very recent). To some extent, plants are

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fire-resistant and capable of re-sprouting from the base. In places where the sphagnum
moss has been totally burnt-out there is evidence of individuals producing wide-spreading
layering systems, forming plants 2-3 m across.
Fitzroya cupressoides is a slow-growing conifer which is capable of reaching up
to 70 m with a girth of four metres and can live longer than 3,600 years (Lara and
Villalba, 1993). In Chile it occurs discontinuously in the coastal range from 39°50′ to
42°30′ S and from 41° to 43°30′ S in the Andes and covers an estimated area of 250,000
ha. Within its range it has an altitudinal range of between 8-1,500 m with a mean annual
precipitation ranging from 2,000 to in excess of 4,000 mm (Villalba, 1990).
During the 18th and 19th centuries, the colonising Spaniards and European
immigrants cleared vast areas of primary forest. This clearance, which was mainly for
agricultural purposes, had a devastating effect on Chile’s most valuable timber tree -
Fitzroya cupressoides. For instance, by the turn of the 19th century, Fitzroya became
commercially extinct on the island of Chiloé; such was the scale of destruction that it is
considered to be one of the most dramatic since the last glaciation (Armesto et al., 1995).
In the Central Depression, north of Puerto Montt, Fitzroya forest may have covered 7,500
ha (Fraver et al., 1999). The huge stumps which can be seen adjacent to Route 5, north of
Puerto Montt, are evidence of how large some of these trees must have been. All that
remains today are 13 remnant forests, five of which contain small forest stands, and eight
with scattered small trees and saplings (Fraver et al., 1999). Since 1975 Fitzroya has been
listed on Appendix I of the Convention on International Trade in Endangered Species
(CITES) and in 1977 the Chilean government declared living Fitzroya a ‘national
monument’ which made it illegal to harvest (see also Lara et al., this volume).

THE PODOCARPACEAE
In the Andes Prumnopitys andina has a distribution range between 35°50′ to
39°30′S (Marticorena and Rodríguez, 1995) and just crosses the Andes into Argentina at
38°S (Tortorelli, 1956). There is also a little known population in the Cordillera de
Nahuelbuta close to Angol (37º50′S 72°50′W) which has, in recent years been much
reduced in size through land conversion to commercial forestry (pers. obs.). Prumnopitys
andina tolerates a wide range of contrasting climates including long dry summers and low
winter temperatures. It is associated with Austrocedrus and small trees and shrubs which
form part of the sclerophyllous vegetation. Nursery grown containerised plants collected
from Nitrao in the Alto Bío Bío (37°41′S) at an altitude of c.1,000 m experienced winter
temperatures of −18°C (Brownless, pers. commun.) and survived with only die-back of
the current year’s late-summer growth. Prumnopitys andina is a predominantly dioecious
species which can attain a height of 15 m and grows between 500 to 1,000 m in altitude
(Rodríguez et al., 1983). Germination trials of the New Zealand Prumnopitys ferruginea
have concluded that seeds continue to mature after seed-fall and can take as long as four
years to germinate (Clout and Tilley, 1992). Similar conclusions for Prumnopitys andina
have been made as a result of propagation trials at the Royal Botanic Garden Edinburgh.
Typically this species produces copious amounts of seed in the wild which accumulates
on the ground beneath the trees only to be eaten by grazing animals such as goats. This
has had a severe impact on the regeneration of this species.
Podocarpus salignus is a dioecious species which occurs in both the Andes and
the coastal cordilleras between 35°50′ to 40°35′S (Marticorena and Rodríguez, 1995). Its
altitudinal range is from 120 m to 1000 m. In the Mediterranean region it grows close to
water courses (where it is sometimes associated with Austrocedrus) but in the same
region the manihuales are well known. These are pure stands of P. salignus which grow
above 1000 m in wet areas not necessarily associated with the margins of water courses
(Donoso, 1981). Due to logging, large trees of Podocarpus salignus are rarely seen but in
primary rainforests which have escaped disturbance trees may reach a height of 20 m.
One such location with large specimens is Cerro Adencul, Traiguén (38°14′S) where they
grow with Austrocedrus chilensis, Cryptocarya alba, Maytenus boaria, Nothofagus
nervosa, N. obliqua, Sophora microphylla, Quillaja saponaria and five species of myrtle

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(pers. obs.). Regeneration is good throughout its natural range with seedlings tolerating a
considerable amount of shade; this is especially supported by copious amounts of
seedlings growing beneath large spreading trees in cultivation in two gardens in SW
England at Caerhayes Castle and Tregothnan.
Saxegothaea conspicua is widely distributed from latitude 36°00′ to 45°45′S. In
northern latitudes (north of 38ºS), where the Mediterranean climate predominates, S.
conspicua is very scarce due to the lack of available water. In its southern range,
particularly in the more mesic evergreen or Valdivian rainforests, it is very abundant. It is
also frequent in the south-central Andes, where it is associated with Nothofagus dombeyi,
N. nervosa and Laureliopsis philippiana, at intermediate altitudes (Donoso et al., 1986;
Donoso, 1993). Where ranges overlap it is often associated with Podocarpus nubigenus.
The latter has a much more extensive north-south distribution than Saxegothaea, from
39°50′ to 50°23′S. Both species have an altitudinal range from sea-level to 1,000 m. In
the coastal range both species occur at low altitudes on poorly-drained marine and fluvio-
glacial deposits and at mid to high elevations on shallow soils developed from mica-
schists (Lusk, 1996). However, it is not commonly found in the Andes in forests that are
dominated by Nothofagus dombeyi, N. nervosa, Laureliopsis philippiana and
Saxegothaea conspicua (Donoso, 1981). Their main tree associates include Amomyrtus
luma, A. meli, Drimys winteri, Eucryphia cordifolia, Dasyphyllum diacanthoides,
Fitzroya cupressoides, Laurelia philippiana, Nothofagus nitida and Weinmannia
trichosperma. Both species are components of the humid forests where they are capable
of attaining heights of 20-25 m. Of all the Chilean conifers, these two species are perhaps
more shade tolerant than any of the others, but the more heavily-shaded parts of the forest
are better exploited by flowering plants (Lusk, 1996).
Lepidothamnus fonkii is the smallest of all the Chilean conifers, forming very
dense, spreading mats not exceeding 25 cm (Pisano, 1983). It occurs discontinuously
from 40°00′ to 55°00′S (Moore, 1983) where it is locally abundant in Patagonian coastal
bogs and coastal mountains. It is most abundant in the magellanic moorlands from 47°00′
to 52°00′S (Marticorena and Rodríguez, 1995) where the rainfall can vary between 1,500
to 3,000 mm (Pisano, 1983). Its common woody associates include Pilgerodendron
uviferum, Empetrum rubrum, Berberis ilicifolia, Berberis buxifolia and Nothofagus
betuloides. At its most northerly location in Parque Nacional Monumento Alerce Costero,
where it grows at an altitude of c. 800 m, it also occurs in bogs with very similar woody
and herbaceous associates.

THE ARAUCARIACEAE
Araucaria araucana has a relatively restricted natural distribution from 37°20′ to
40°20′S. This is thought to be a remnant of a much bigger distribution before the logging
activities associated with European colonisation (Veblen and Delmastro, 1976). Its main
Chilean distribution is in the Andes where it has an altitudinal range from 900 to 1,800 m
on layers of volcanic ash deposits or soils derived from metamorphic and sedimentary
rocks. On the western part of the range the annual precipitation reaches 4,000 mm
dropping to 1,100 mm on the eastern side in Argentina, where typically the forests
experience dry summers and high winter snowfalls. At the timber-line, Araucaria forms
an open canopy with Nothofagus pumilio as the understorey; at lower altitudes it is
associated with N. dombeyi. In Chile there is a disjunction in the distribution of A.
araucana as it also occurs on the coast in the Cordillera de Nahuelbuta, west of Angol
(37°20′ and 38°40′S). These forests, which range from 600 to 1,400 m (Veblen, 1995),
are on soils which have derived from ancient granitic and metamorphic rocks with an
annual rainfall of between 1,500-2,500 mm. The main woody associated species include
Nothofagus antarctica, N. nervosa, N. obliqua, Pseudopanax laetevirens, Desfontainia
spinosa and Drimys andina. A. araucana is predominantly dioecious and capable of
growing to 45 m tall and living for over 1,300 years. It is well adapted to fire and has
thick, plate-like bark and epicormic buds which sprout following the occurrence of fire.
Seed is eaten, and perhaps dispersed, by a variety of animals and birds, including rodents

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and parakeets. Domestic animals and man have also played a significant role and some of
the more outlying trees to the present distribution may be attributed to native Mapuche
people.

CONCLUSION
All nine native Chilean conifers are listed by the World Conservation Union
(IUCN) as threatened (Farjon and Page, 1999). Araucaria araucana and Fitzroya
cupressoides are listed as Chilean Natural Monuments, which prohibit their logging and
they are also listed on Appendix I under the Convention on International Trade in
Endangered Species (CITES), which prevents international trade. The rainforests of Chile
continue to be under tremendous pressures due to their non-sustainable use. Such
activities as burning and logging of native forests in order to make way for extensive
plantations of non-native species such as Pinus radiata and Eucalyptus globulus, will
continue to have a knock-on effect on the distribution and ecology of Chilean conifers.

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