Woodland communities in the Chilean cold-temperate zone (Baker and Pascua basins): Floristic composition and morpho-ecological transition

Abstract

This study describes the floristic composition and morpho-ecological transition of woodlands along a climatic gradient in the southern cold temperate zone of Chilean Patagonia. A total of 256 phytosociological relevés were performed across a 150 km NE-SW transect to record vascular plant species. Classification (cluster analysis) and ordination (principal component analysis) techniques were used to segregate and examine the communities. Biodiversity indicators including richness and abundances of species natives and exotics, importance values, Raunkiaer plant life-forms, diversity indices and indicator species were calculated to describe community attributes. Beta diversity was analysed using the Jaccard index. Additionally, the current anthropogenic disturbances affecting this vegetation are discussed. In total, 11 woodland communities belonging to 3 morpho-ecological groups were segregated: a) meso-hygromorphic woodlands belonging to the Baker basin, mostly composed of deciduous forests containing relatively moderate values of richness and diversity but high richness of exotics, b) hygromorphic woodlands belonging to the southern segment of the Baker basin and along the Pascua basin, composed of evergreen forest containing the relatively highest values of richness and diversity and very low richness of exotics and c) high-Andean dwarf woodlands distributed at high elevations in both basins, composed of deciduous krummholz containing the lowest richness and diversity and no exotics. The replacement of deciduous by evergreen communities at low elevations occurs around the latitude 48°S. Anthropogenic disturbances like logging by rural landowners, overgrazing by livestock and road construction are promoting biological invasions in the Baker basin forests, while the forests in the Pascua basin remain pristine since no human population occurs there.

Full text

141
Woodland communities in the Chilean cold-temperate zone (Baker
and Pascua basins): Floristic composition and morpho-ecological
transition
Comunidades leñosas en la zona chilena frío-templada (cuencas de los ríos Baker y
Pascua): Composición florística y transición morfo-ecológica
OSVALDO J. VIDAL1, JAN R. BANNISTER1, VÍCTOR SANDOVAL2, YESSICA PÉREZ3 & CARLOS RAMÍREZ4
1Institute of Silviculture, Faculty of Forest and Environmental Sciences, University of Freiburg, Tennenbacherstrasse 4, D-79106,
Freiburg im Breisgau, Germany.
2Instituto de Manejo Forestal, Facultad de Ciencias Forestales y Recursos Naturales, Universidad Austral de Chile, Casilla 567,
Isla Teja, Valdivia. Chile.
3Instituto de Botánica, Facultad de Ciencias, Universidad Austral de Chile, Casilla 567, Isla Teja, Valdivia. Chile.
4Departamento de Ecología, Facultad de Ciencias Biologicas, Universidad Católica de Chile, Alameda 340, Santiago, Chile.
osvaldo.vidal@waldbau.uni-freiburg.de
ABSTRACT
This study describes the floristic composition and morpho-ecological transition of woodlands along a climatic gradient in
the southern cold temperate zone of Chilean Patagonia. A total of 256 phytosociological relevés were performed across
a 150 km NE-SW transect to record vascular plant species. Classification (cluster analysis) and ordination (principal
component analysis) techniques were used to segregate and examine the communities. Biodiversity indicators including
richness and abundances of species natives and exotics, importance values, Raunkiaer plant life-forms, diversity indices
and indicator species were calculated to describe community attributes. Beta diversity was analysed using the Jaccard
index. Additionally, the current anthropogenic disturbances affecting this vegetation are discussed. In total, 11 woodland
communities belonging to 3 morpho-ecological groups were segregated: a) meso-hygromorphic woodlands belonging to
the Baker basin, mostly composed of deciduous forests containing relatively moderate values of richness and diversity
but high richness of exotics, b) hygromorphic woodlands belonging to the southern segment of the Baker basin and along
the Pascua basin, composed of evergreen forest containing the relatively highest values of richness and diversity and very
low richness of exotics and c) high-Andean dwarf woodlands distributed at high elevations in both basins, composed
of deciduous krummholz containing the lowest richness and diversity and no exotics. The replacement of deciduous by
evergreen communities at low elevations occurs around the latitude 48°S. Anthropogenic disturbances like logging by rural
landowners, overgrazing by livestock and road construction are promoting biological invasions in the Baker basin forests,
while the forests in the Pascua basin remain pristine since no human population occurs there.
KEYWORDS: Biogeographical transition, southern cold temperate zone, anthropogenic disturbances, pristine forests, Aysén.
RESUMEN
Este estudio describe la composición florística y la transición morfo-ecológica de las comunidades leñosas ocurriendo
a través de un gradiente climático en la zona templada fría de la Patagonia chilena. Se establecieron un total de 256
relevamientos fitosociológicos a través de un transecto NE-SO de 150 km para registrar las especies de plantas vasculares.
Técnicas de clasificación (análisis de conglomerados) y ordenación (análisis de componentes principales) fueron usadas
para segregar y examinar comunidades. Se computaron indicadores de biodiversidad incluyendo riqueza y abundancia
de especies nativas y exóticas, valores de importancia, formas de vida de Raunkiaer, índices de diversidad y especies
indicadoras para describir atributos comunitarios. La diversidad Beta fue analizada usando el coeficiente de Jaccard. Se
discuten también las perturbaciones antropogénicas que actualmente afectan a la vegetación. En total se segregaron 11
comunidades pertenecientes a tres grupos ecológicos: a) comunidades leñosas meso-higromórficas pertenecientes a la
cuenca del Baker, conformada principalmente de bosques caducifolios conteniendo valores relativos intermedios de riqueza
y diversidad, pero las mayores riquezas de exóticas; b) comunidades leñosas higromórficas pertenecientes al segmento sur
de la cuenca del río Baker y a través de toda la cuenca del río Pascua, compuesta de bosques siempreverdes conteniendo
los mayores valores de riqueza y diversidad y muy baja riqueza de exóticas, y c) comunidades leñosas achaparradas alto-
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Gayana Bot. 68(2), 2011
andinas, distribuidas en lugares de alta elevación en ambas cuencas, compuestas de krummholz conteniendo la menor
riqueza y diversidad, sin presencia de especies introducidas. El reemplazo de comunidades caduficolias por siempreverdes
en sentido norte-sur sucede alrededor de la latitud 48°S. Perturbaciones antrópicas como la tala de madera por propietarios
rurales, sobrepastoreo por ganado doméstico y ampliaciones en los caminos, están provocando invasiones biológicas en
los bosques de la cuenca del río Baker, mientras que los bosques de la cuenca del río Pascua, donde no ocurre poblamiento
humano, permanecen prístinos.
PALABRAS CLAVE: Transición biogeográfica, zona templada fría, perturbaciones antrópicas, bosques prístinos, Aysén.
INTRODUCTION
The Baker and Pascua rivers are located in the Aysén Region
in northern Chilean Patagonia. The former is about 175 km
in length, originating at the southern tip of Bertrán Lake and
ending in the Baker Channel. The Pascua River, in turn, is 67
km in length, originating at the northwest side of O’Higgins
Lake and ending at the Calén Fjord. Adjacent to both rivers
is an extensive 150 km long basin-strip system, distributed
along a steep precipitation gradient with annual precipitation
ranging from approximately 350 mm in the continental
areas to 3,000 mm at the archipelagic locations (Amigo &
Ramírez 1998). These basins are flanked to the northwest
by the North Patagonian Ice Fields and to the southwest by
the archipelagic zone. The entire area is transected by the
Andean mountain range, with elevations ranging from sea
level to 1,200 m. The area is thus a tangled corridor where
floristic elements transit latitudinally and altitudinally
throughout the “Southern Cold Temperate Zone” (Du Rietz
1960, Holdgate 1960) (Fig. 1). The vegetation in this area
is diverse and includes Patagonian steppes, shrublands,
deciduous forests, perennial forests at low elevations and
moorland vegetation above the timberline (e.g. Hueck 1978,
Veblen & Schlegel 1982, Luebert & Pliscoff 2006). Given
the difficult access to some parts of the region and its low
human population density (0.8 inhabitants per km2), the
region contains landscapes in completely pristine conditions
(Martínez-Harms & Gajardo 1998).
The first descriptions of the area’s vegetation correspond
to phytogeographical studies made at the beginning of the
20th century by Reiche (1934) and Hambleton (1936),
who described the forest vegetation and drafted some
geographical limits. Hambleton (1936) mentioned the
forests dominated by Nothofagus dombeyi (Mirb.) Oerst
distributed in the northern area of the Baker basin. Reiche
(1934), meanwhile, mentioned the Nothofagus pumilio
(Poepp. et Endl.) Krasser forests in the central area of the
Baker basin and the presence of steppe-like shrublands in
the northeastern area. He described forests in the southern
area as being dominated by Nothofagus nitida (Phil.) Krasser
and krummholz communities and ‘carpet vegetation’ above
the timberline. Godley (1960) established latitude 48°S as
the limit where deciduous forests are replaced by perennial
forests and subantarctic moorlands.
Although later studies have described and delineated the
woodland vegetation of the area (e.g. Fuenzalida & Pisano
1967, Pisano 1972, Hueck 1978, Veblen & Schlegel 1982,
Luebert & Pliscoff 2006), none have been able to fully
compile its complex structure and diversity, partly due to
the difficult access of some of these places. Moreover, some
of the authors considered just dominant species, omitting
valuable information related to target non-dominant
ones. Detailed information can be not only interesting for
biogeographers looking to explain biotic patterns, but also
useful for managers requiring quantified information to take
management decisions (Noss 1990). In this context, this study
aimed: 1) to classify and describe the woodland communities
along the Baker and Pascua river basins, 2) to compare the
floristic diversity among these communities, 3) to delineate
the distribution of native woodland communities and 4) to
describe human disturbances affecting this vegetation.
MATERIALS AND METHODS
STUDY AREA
The study was carried out in the Baker and Pascua basins,
located in the administrative Region of Aysén in northern
Chilean Patagonia (Fig. 1). This area (∼150,000 ha) lies
approximately between 47°03’S and 48°21’S and 73°19’
W and 72°27’ W. The altitudinal range is from sea level
to ∼1,200 m. Meteorological data from the study area are
not available, but a notable NE-SW climatic gradient can
be observed. Annual temperatures recorded at the nearest
meteorological stations range from 10°C (Chile Chico,
46º36'S - 71º43'W) to 7.1ºC (Puerto Edén, 49º08' S -
74º25'W), while precipitation ranges from 355 mm (Chile
Chico) to 3,033 mm (Puerto Edén). These two locations
belong to the dry and ultraperhumid bioclimatic belts,
respectively, in the temperate zone of Chile (Amigo &
Ramírez 1998). The vegetation in the study area includes
arid steppes, xerophylous shrublands and deciduous forests
in NE locations, being gradually replaced toward the SW by
hygrophilous vegetation such as peatlands, shrublands, and
perennial forests (Luebert & Pliscoff 2006).
FIELD METHODS AND TAXONOMICAL DETERMINATIONS
Between April 2006 and March 2007, four field expeditions
were organized in order to cover the entire ∼150 km NE-
SW transect in the study area. Sample sites were selected

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Woodlands in the Chilean cold-temperate zone: VIDAL, O. ET AL.
according to a stratified design using aerial photographs
and were accessed by means of horses, vehicles, boats and
helicopters because of the difficulty of the terrain.
In each site a 500 m2 phytosociological relevé
(Mueller-Dombois & Ellenberg 1974) was delimited to
record all vascular plants and to estimate the percentage
of abundances of each. Topographical features such as
elevation, slope inclination, slope aspect and GPS location
were also recorded. Voucher specimens were collected and
determined using taxonomical literature (Cabrera 1949,
Muñoz-Schick 1980, Moore 1982, Matthei 1995, Ruiz 2001,
Landrum 2003, Rodríguez & Quezada 2003, Marticorena
2006). Nomenclature follows Henríquez et al. (1995) and
Rodríguez et al. (2008).
DATA ANALYSIS
We constructed a relevé-by-species abundance matrix
containing a total of 256 relevés and 201 species. Species
abundance data were transformed by using the logarithmic
transformation (log (x+1)) to get an approximation close
to normal and to stabilize the variance (McCune & Grace
2002), as required for multivariate methods. In order to
detect plant communities, the relevé-by-species abundance
matrix was classified by performing hierarchical
agglomerative clustering using Ward’s minimum variance
criterion (Euclidean distance) to minimize the increase
in within-group variance (Hammer et al. 2001). Sites
belonging to a particular cluster-group (i.e. community)
were pooled and the new community-by-species matrix
was ordinated using principal component analysis (PCA)
(Euclidean distance) to examine the structure of nodal
communities. From each community-by-species matrix
the structural and compositional attributes of vegetation
biodiversity were extracted (Noss 1990), i.e. mean
species richness, cumulative abundances, importance
values (Wikum & Shanholtzer 1978), diversity indices
(Shannon and Berger-Parker, Moreno 2001), Raunkiaer
plant life-forms (Mueller-Dombois & Ellenberg 1974) and
exotic-native species ratio (Noss 1990). Indicator species
analysis (Dufrêne & Legendre 1997) was performed to
assess fidelity of particular species to each community as
diagnostic species. In order to compare diversity between
communities, a beta diversity comparison matrix using the
Jaccard coefficient (Lapin & Barnes 1995) was performed
from the pooled database. Finally the patterns of biological
invasions were explored by evaluating correlation between
elevation of sites, distance to roads and number of exotic
plant species. Structural and compositional attributes of
communities, cluster analysis, PCA and correlations were
performed using the PAST statistical package version 1.91
(Hammer et al. 2001). ISA was performed using the labdsv
indval Package for R statistical software.
FIGURE 1. Map of Aysén, northern Chilean Patagonia, showing the study area.
FIGURA 1. Mapa de Aysén, Norte de Patagonia Chilena, mostrando el área de estudio.

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Gayana Bot. 68(2), 2011
RESULTS
CLASSIFICATION, ORDINATION AND DESCRIPTION OF COMMUNITIES
Nine woodland communities were classified after clustering
(Fig. 2), but 2 sub-groups, krummholz dominated by
Nothofagus pumilio (5 relevés) and krummholz dominated
by Nothofagus antarctica (G.Forst.) Oerst. (5 relevés), were
arbitrarily reassigned as communities based on their habitat
features (lower canopy heights), lower number of species
and site topography (high elevation sites, steep slopes).
The ordination diagram (Fig. 3) showed the nodal
communities distributed in 3 groups along the two principal
components, explaining 53.8% of the variance. These
components can be hypothesized as environmental factors
affecting the distribution of plant communities. We thus
hypothesized that component 1 (x-axis) was temperature
and component 2 (y-axis) was precipitation. The first group
contained 3 communities distributed in colder and more
humid places and included forests dominated by Nothofagus
betuloides (Mirbel) Oersted, forests dominated by
Nothofagus nitida and forests dominated by Pilgerodendron
uviferum (D.Don) Florin. The second group contained 5
communities distributed in warmer places with intermediate
levels of precipitation and included Nothofagus antarctica
forests, Embothrium coccineum shrublands, Nothofagus
antarctica shrublands, Nothofagus dombeyi forests and
Nothofagus pumilio forests. The third group of communities
can be considered to be distributed in dry and cold places and
included krummholz dominated by Nothofagus antarctica
and krummholz dominated by Nothofagus pumilio. A brief
description of the plant communities is provided below, based
on dominant species, importance values (IV), diagnostic
species identified by indicator species analysis (ISV)
(Annex 1), plant life-forms (Fig. 4), and cumulative richness
of native and exotic species. Communities were named
according to the dominant species, but a phytosociological
name from the literature was also provided:
1) Nothofagus nitida forests (Luzuriago polyphyllae-
Nothofagetum nitidae). Evergreen and multi-layered forests
located mainly in lowland and flat areas (0 - 400 m) close to
river courses and lakes. Trees reach up to a height of 30 m.
The vegetation structure is complex, dominated in the upper
layer by Nothofagus nitida (IV 32.2) but co-dominated by
other phanerophytes like Podocarpus nubigena Lindl. (IV
18.7), Drimys winteri J.R.Forst. et G.Forst. (IV 16.5) and
Desfontainia fulgens D.Don (IV 3.7), among others. There is
a notable presence of plants growing as epiphytes, including
Hymenophyllum krauseanum Phil. (IV 18.5), Philesia
magellanica J.F.Gmel. (IV 5.3) and Mitraria coccinea
Cav. (IV 5.0). The shrub layer is also rich, containing
microphanerophytes like Pseudopanax laetevirens (Gay)
Franchet (IV 10.5) and Chusquea culeou E. Desv. (IV 4.0).
On the ground are hemicryptophytes like Blechnum penna-
marina (Poir.) Kuhn (IV 9.3), Blechnum magellanicum
(Desv.) Mett. (IV 7.8) and Rubus geoides Sm. (IV 3.9), among
many other species. Exotic elements are not important but
include some species like Holcus lanatus L.(IV 0.6), Prunella
vulgaris L. (IV 0.6) and Trifolium repens L. (IV 0.5), among
others. Diagnostic species with the greatest ISA values were
Nothofagus nitida (ISA 1.0), Hymenophyllum krauseanum
(ISV 0.7) and Griselinia rusciflora (Clos) Ball. (ISV 0.56).
FIGURE 2. Dendrogram showing floristic relationships among the communities segregated in the present study. The arrow on the y-axis
indicates the segregation point. Communities 5 and 10 were arbitrarily segregated (see results section).
FIGURA 2. Dendrograma mostrando las relaciones florísticas entre las comunidades segregadas en el presente estudio. La flecha en el eje y
muestra el punto de segregación. Las comunidades 5 y 10 fueron segregadas arbitrariamente (ver la sección de resultados).

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Woodlands in the Chilean cold-temperate zone: VIDAL, O. ET AL.
No human disturbance affected these stands, although a
few exotic species were detected. We recorded 67 species
belonging to these stands, 7 of them exotic. These forests
were distributed in the central and southern section of the
Baker basin and in the northern section of the Pascua basin.
2) Pilgerodendron uviferum forests (Empetro-
Pilgerodendronetum uviferae). Evergreen resinous multi-
layered forests associated with flat and inundated areas (0
- 400 m) where extensive cover of the moss Sphagnum
magellanicum Brid on the ground was observed. Dominant
trees reached up to 6 m in height. Pilgerodendron uviferum
dominated in cover and frequency (IV 30.5) at the upper
layer, and Empetrum rubrum Vahl ex Willd. (IV 26.4)
co-dominated at the ground layer. Other important
species were Gaultheria mucronata (L.f.) Hook. et
Arn. (IV 17.7), the epiphyte Philesia magellanica (IV
12.4) and the phanerophyte Nothofagus antarctica (IV
8.8). Diagnostic species with the greatest ISV values
included Pilgerodendron uviferum (ISV 0.79), Oreobolus
obtusangulus Gaudich. (ISV 0.57) and Empetrum rubrum
(ISV 0.49). Large areas of these forests were burned
during colonial times and remained standing, some of
them providing seed trees. We recorded 36 species in these
stands, all of them native. The stands were distributed in
the southern segment of the Baker basin.
3) Nothofagus betuloides forests (Nothofagetum
betuloidis). Evergreen forests associated with gentle
slopes and mountainous areas from sea level to ∼600
m elevation. Vegetation structure was multi-layered,
dominated by Nothofagus betuloides (IV 30.9) in the
upper tree layer, but co-dominated by other trees like
Desfontainia fulgens (IV 10.7), Podocarpus nubigena
(IV 10.4) and Drimys winteri (IV 9.5). The presence
of many epiphytes was detected on the trees, including
Philesia magellanica (IV 14.6) and ferns like Serpyllopsis
caespitosa (Gaudich.) C.Chr. (IV 6.0), Hymenophyllum
tortuosum Hook. et Grev. (IV 2.2) and Hymenophyllum
pectinatum Cav. (IV 1.1). The shrub layer was also rich
in species and included many microphanerophytes like
Gaultheria mucronata (IV 17.2), Berberis ilicifolia L.f.
(IV 8.0) and Lebetanthus myrsinites (Lam.) Dusen (IV
7.8), among others. Ground vegetation was also rich in
species and includes Blechnum penna-marina (IV 5.7),
Sticherus quadripartitus (Poir.) Ching (IV 5.2) and
Blechnum magellanicum (IV 4.7). Diagnostic species
with the greatest ISV values included Nothofagus
FIGURE 3. Diagram of principal component analysis (PCA) summarizing ordination of native woodland communities in the Baker and Pascua
basins. The components explain 53.8% of the variance. Symbols: = meso-hygromorphic woodlands, mostly deciduous woodlands, =
hygromorphic woodlands, perennial woodlands, and Δ= high-Andean dwarf woodlands, krummholz communities.
FIGURA 3. Diagrama basado en el análisis de componentes principales (PCA) resumiendo la ordenación de las comunidades leñosas nativas
en las cuencas del Baker y del Pascua. Los componentes explican el 53,8% de la varianza. Símbolos: = comunidades leñosas meso-
higromórficas, principalmente conformadas por bosques caducifolios, = comunidades leñosas higromórficas, conformadas por bosques
perennes y Δ = matorrales achaparrados alto-andinos, comunidades del tipo krummholz.

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Gayana Bot. 68(2), 2011
FIGURE 4. Raunkiaer life-form spectrum of different woodland communities in the Baker and Pascua basins according to species number
(relative richness) and species abundances (relative cover) of species. Raunkiaer plant life-forms: Pha= Phanerophyte, Epi= Epiphytes,
Cha= Chamaephyte, Hem= Hemicryptophyte, Geo= Geophyte, The= Therophyte. Name of communities: 1. Nothofagus nitida forests, 2.
Pilgerodendron uviferum forests, 3. Nothofagus betuloides forests, 4. Nothofagus pumilio forests, 5. Nothofagus pumilio krummholz, 6.
Nothofagus dombeyi forests, 7. Embothrium coccineum shrublands, 8. Nothofagus antarctica forests, 9. Nothofagus antarctica shrublands,
10. Nothofagus antarctica krummholz, 11. Pinus contorta plantations.
FIGURA 4. Espectros biológicos basados en las formas de vida de Raunkiaer para las diferentes comunidades leñosas en las cuencas de
los ríos Baker y Pascua, de acuerdo al número de especies (riqueza relativa) y abundancia de especies (cobertura relativa). Formas de
vida de Raunkiaer: Pha= fanerófitos, Epi= epífitos, Cha= caméfitos, Hem= hemicriptófitos, Geo= geófitos, The= terófitos. Nombre de
las comunidades: 1. Bosques de Nothofagus nitida, 2. Bosques de Pilgerodendron uviferum, 3. Bosques de Nothofagus betuloides, 4.
Bosques de Nothofagus pumilio, 5. Krummholz de Nothofagus pumilio, 6. Bosques de Nothofagus dombeyi, 7. Matorrales de Embothrium
coccineum, 8. Bosques de Nothofagus antarctica, 9. Matorrales de Nothofagus antarctica, 10. Krummholz de Nothofagus antarctica, 11.
Plantación de Pinus contorta.
betuloides (ISV 0.7), Serpyllopsis caespitosa (ISV 0.5)
and Desfontainia fulgens (ISV 0.38), among others. No
human disturbances were detected in these stands. We
recorded a total of 68 species, all of them native. These
stands were detected in the southern section of Baker
and Pascua basins.
4) Nothofagus pumilio forests (Mayteno-Nothofagetum
pumilionis). Deciduous forests associated with gentle slopes
from ∼100 m to ∼1,000 m in elevation. Dominant trees
reached up to 30 m in height and the vegetation structure was
3-layered, with Nothofagus pumilio (IV 52.2) dominating at
the upper layer. Gaultheria mucronata (IV 17.7), Maytenus
disticha (Hook.f.) Urban (IV 7.5), Berberis microphylla
Hort ex. K.Koch (IV 6.3) and Berberis darwinii Hook. (IV
6.3) were the most important microphanerophytes in the
shrub layer. The ground layer was rich in species, with many
hemicryptophytes like Blechnum penna-marina (IV 7.8),
Fragaria chiloensis (L.) Duch. (IV 7.2) and Adenocaulon
chilense Poepp. ex Less. (IV 6.7) as natives, and Trifolium
repens L. (IV 4.1) and Taraxacum officinale (L.) Weber
(IV 3.2) as the most important exotics. Diagnostic species
included Nothofagus pumilio (ISV 0.55), Viola maculata
Cav. (ISV 0.49), Maytenus disticha (ISV 0.43), Adenocaulon
chilense (ISV 0.25) and the orchid Codonorchis lesonii
(Brongn.) Lindl. (ISV 0.18). The stands were affected
by human disturbances like the use of wood for fuel and
overgrazing by livestock. A consequent modification of
the structure (e.g. reduction in canopy cover) and floristic
composition of these forests (particularly the introduction
of exotic species) was observed. We recorded a total of 79
species in these stands, 19 of them exotic. The distribution
of these forests was restricted to the Baker basin.
5) Nothofagus pumilio krummholz (Empetro-
Nothofagetum pumiliae). Deciduous and dense shrubland
reaching up to 2 m in height, poor in species and associated
with high elevation areas at the timberline from ∼1,000 to
1,200 m. The vegetation structure was reduced to only 2
strata, with the upper layer dominated by N. pumilio (IV
116.6) and co-dominated by Escallonia alpina Poepp. &
Endl. (IV 4.9). The floristic composition was poor, with
Empetrum rubrum (IV 31.7) and Gaultheria pumila (L.f.)
D.J.Middleton (IV 20.3) as chamaephytes and some
hemicryptophytes like Rubus geoides (IV 21.6) and Luzula
racemosa (IV 4.9). Diagnostic species included Gaultheria
pumila (ISV 0.34) and Rubus geoides (ISV 0.29). Stands
were pristine and not affected by human disturbances due
to their difficult accessibility. We recorded a total of 6
species, all of them native. The distribution of these stands
was restricted to the Baker basin.

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6) Nothofagus dombeyi forests (Chrysosplenio-
Nothofagetum dombeyi). Monospecific perennial forests
located in low elevation areas near water bodies from sea
level to ∼500 m. Trees reached up to 40 m in height. The
vegetation structure was 3-layered, with N. dombeyi (IV
64.8) as the dominant species and Gaultheria mucronata
(IV 15.0), Berberis microphylla (IV 7.2), Embothrium
coccineum J.R.Forst. et G.Forst (IV 6.6), and Berberis
darwinii (IV 6.4) as important microphanerophytes in the
shrub layer. The ground vegetation was rich in species, with
Blechnum penna-marina (IV 12.6), Adenocaulon chilense
(IV 6.5) and Acaena ovalifolia Ruiz & Pav. (IV 6.0) being
the most important native hemicryptophytes. Some exotics
included Taraxacum officinale (IV 3.2), Holcus lanatus (IV
2.0) and Hypochaeris radicata L. (IV 1.4), among others.
Diagnostic species included Nothofagus dombeyi (ISV
0.89), Ribes magellanicum Poir. (ISV 0.18) and Viola reichei
Skottsb. (ISV 0.16). Stands are affected by logging due to
the use of wood as fuel by the rural landowners. Overgrazing
of livestock was also observed in these stands, with evident
browsing of shrubs and regenerating trees. We recorded a
total of 81 species, 17 of them exotic. The distribution of
these stands was restricted to the Baker basin.
7) Embothrium coccineum shrublands (Embothrio-
Pernettietum mucronatae). Deciduous tree shrublands
colonizing gentle slopes fluctuating from sea level to ∼900
m. The vegetation structure was 3-layered, with Embothrium
coccineum as the dominant phanerophyte but which did not
form a continuous canopy (IV 33.3). Gaultheria mucronata
was the dominant microphanerophyte at the shrub layer (IV
44.0). Other phanerophyte species were Escallonia rosea
Griseb. (8.7), Berberis microphylla (IV 7.0) and Nothofagus
antarctica (IV 5.3). The ground layer was rich in species,
containing some chamaephytes like Baccharis magellanica
(Lam.) Pers. (IV 14.4) and Empetrum rubrum (IV 12.0),
and hemicryptophytes like Festuca pyrogea Speg. (IV 5.0),
Blechnum penna-marina (IV 5.0) and Fragaria chiloensis
(IV 4.4). Important exotics were Hypochaeris radicata (IV
2.5) and Rumex acetosella L. (IV 2.1). Diagnostic species
included Embothrium coccineum (ISV 0.6), Escallonia
rosea (ISV 0.51) and Baccharis magellanica (ISV 0.49)
as phanerophytes. Lycopodium paniculatum Desv. (ISV
0.39) and Gavilea odoratissima Poepp. (ISV 0.34) were
included as hemicryptophyte and geophyte diagnostic
species, respectively. These shrublands were affected to a
small degree by human disturbances derived from livestock
production. We recorded a total of 59 species, 8 of them
exotic. The distribution of these shrublands was restricted to
the Baker basin, especially in the southern section.
8) Nothofagus antarctica forests (Chusqueo-Nothofagetum
antarcticae). Deciduous forests associated with areas of
depressions in the relief, close to river meanders from sea
level to ∼80 m. The structure was 3-layered, with Nothofagus
antarctica (IV 47.0) as the dominant species, reaching up to
15 m in height. Chusquea culeou (IV 32.4) and Embothrium
coccineum (IV 5.8) were the co-dominant species in the
upper and shrub layer. Microphanerophytes in the shrub
layer included Escallonia virgata (IV 13.8), Gaultheria
mucronata (IV 9.2) and Berberis microphylla (IV 8.2). The
ground vegetation was rich in species, containing many
hemicryptophytes like Blechnum penna-marina (IV 16.6),
Acaena ovalifolia (IV 7.3), and Rubus geoides (IV 4.2).
Exotics included Prunella vulgaris (IV 4.4), Taraxacum
officinale (IV 4.2) and Trifolium repens (IV 3.4). Diagnostic
species were Chusquea culeou (ISA 0.76), Escallonia
virgata (ISA 0.40) and Blechnum penna-marina (ISA 0.20),
among others. In these forests we recorded a total of 31
species, 6 of them exotic. The distribution of these stands
was restricted to the southern section of the Baker basin.
9) Nothofagus antarctica shrublands (Anemone
polyphyllae-Nothofagetum antarcticae). Deciduous tree
shrublands reaching up to 8 m in height, located in flat
areas from sea level to ∼500 m. The structure was 3-layered
with Nothofagus antarctica dominating the upper layer (IV
47.2). Other phanerophytes such as Berberis microphylla
(IV 6.2), Discaria chacaye (IV 4.8), and Ribes cucculatum
(IV 3.5) were present but had values of little importance.
The understory was rich in species, containing many natives
like Festuca pyrogea (IV 8.7), Blechnum penna-marina (IV
8.0), Anemone multifida (IV 7.6) and Fragaria chiloensis
(IV 6.6), and many exotics like Taraxacum officinale (IV
6.0), Holcus lanatus (IV 4.1), Trifolium repens (IV 3.6) and
Achillea millefolium (IV 3.5), among others. The diagnostic
species included Anemone multifida (ISV 0.40) and
Discaria chacaye (ISV 0.29) as natives, and some exotics
like Achillea millefolium (ISV 0.26), Cerastium arvense
(0.16) and Carduus nutans (ISV 0.13), among others. Cattle
heavily affected these shrublands because the ground plant
cover was used as forage. Pinus contorta populations have
also invaded some stands. In total we recorded 107 species
in these stands, 38 of them exotic. The distribution of these
forests occurred along the entire Baker basin.
10) Nothofagus antarctica krummholz (Senecio
acanthifoliae-Nothofagetum antarcticae). Deciduous dwarf
shrublands reaching up to a height of 1 m, poor in species and
located in highlands from 600 to 900 m. The structure was 2-
layered and dominated by Nothofagus antarctica (IV 85.9)
in the upper layer as the only phanerophyte. The understory
cover was dominated by Gunnera magellanica (IV 28.1) and
Senecio acanthifolius (IV 23.3) as hemicryptophytes, and
Bolax caespitosa (IV 17.5) and Empetrum rubrum (IV 10.4)
as chamaephytes. The diagnostic species included Senecio
acanthifolius (ISV 0.92), Gunnera magellanica (ISV 0.62)
and Bolax caespitosa (ISV 0.60), among others. No human

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Gayana Bot. 68(2), 2011
disturbances were detected in these stands. In total we
recorded 11 species, all of them native. These stands were
distributed along the entire Pascua basin.
11) Pinus contorta plantations. Exotic monoculture
located near the village of Cochrane, between 200 - 300
m. The structure was 3-layered with Pinus contorta (IV
67.2) having the major importance value. Nothofagus
antarctica (IV 26.3) was also an important phanerophyte.
Microphanerophytes in the shrub layer included Berberis
microphylla (IV 7.3) and Gaultheria mucronata (IV 2.4).
The ground vegetation was poor in species and included
some natives like Nertera granadensis (IV 13.5), Acaena
ovalifolia (IV 7.3) and Osmorhiza chilensis (IV 4.7),
and some exotics like Agrostis stolonifera (IV 44,3),
Taraxacum officinale (IV 12.2) and Trifolium repens (IV
2.4). Diagnostic species were Pinus contorta (ISV 1.0),
Agrostis stolonifera (ISV 0.87) and Nertera granadensis
(ISV 0.51). Some Pinus contorta individuals were growing
outside of plantations and invading native communities.
In total we recorded 13 species, 4 of them exotic. The
distribution of these stands is restricted to close to the
village of Cochrane in the Baker basin.
COMMUNITY COMPARISONS AND DISTURBANCE PATTERNS
Mean cover and richness of native and exotic plant
species as well as diversity indices (Shannon and Beger-
Parker) are presented in Table I. Forest communities
dominated by Nothofagus nitida, Pilgerodendron
uviferum and Nothofagus betuloides have the highest
richness as well as cover and Shannon diversity values,
indicating relatively equitable participation in cover for
many species. In these communities, exotic elements
are virtually absent, with only Nothofagus nitida
forests having low values of exotics. On the other hand,
communities containing the lowest values of richness are
Nothofagus pumilio krummholz, Nothofagus antarctica
krummholz and the Pinus contorta plantations. These
communities also contain the lowest Shannon index
values but have the highest Berger-Parker index values,
indicating the higher cover of the dominant woods. The
communities most invaded by exotics are Nothofagus
antarctica shrublands, Nothofagus pumilio forests and
Pinus contorta plantations, while 4 communities have
no exotics (Pilgerodendron uviferum forests, Nothofagus
betuloides forests, Nothofagus antarctica krummholz and
Nothofagus pumilio krummholz).
Similarities among communities based on the Jaccard
coefficient vary from 0.49 between Nothofagus dombeyi
forests and Embothrium coccineum shrublands (the highest
value) to no similarity (value 0 for the index) between
Nothofagus pumilio krummholz and Pinus contorta
plantations, revealing that each community has its own
identity in terms of species composition (Table II).
TABLE I. Diversity structure of woody communities in the Baker and Pascua basins.
TABLA I. Diversidad estructural de las comunidades leñosas en las cuencas de los ríos Baker y Pascua.
WOODLAND COMMUNITY NATI V E
RICHNESS
EXOTIC
RICHNESS
NATI V E
COVERS
EXOTIC
COVERS
SHANNON
INDEX
BERGER PARKER
INDEX
Nothofagus nitida forests 14.8 ± 3.4 0.4 ± 0.8 140.5 ± 31.5 1.0 ± 4.2 1.878 ± 0.311 0.366 ± 0.122
Pilgerodendron uviferum forests 20.4 ± 3.2 - 173.1 ± 20.6 - 2.175 ± 0.148 0.305 ± 0.058
Nothofagus betuloides forests 16.2 ± 3.9 - 149.7 ± 44.5 - 1.995 ± 0.340 0.313 ± 0.121
Nothofagus pumilio forests 13.8 ± 3.1 2.2 ± 2.0 123.7 ± 34.4 7.3 ± 11.4 1.636 ± 0.348 0.485 ± 0.125
Nothofagus pumilio krummholz 3.5 ± 1.4 - 110.0 ± 14.5 - 0.371 ± 0.329 0.891 ± 0.104
Nothofagus dombeyi forests 11.6 ± 3.5 1.6 ± 2.5 127.0 ± 34.0 4.0 ± 8.0 1.276 ± 0.379 0.601 ± 0.168
Embothrium coccineum shrublands 12.3 ± 2.5 0.8 ± 0.9 132.5 ± 49.1 0.8 ± 0.9 1.609 ± 0.189 0.429 ± 0.110
Nothofagus antarctica forests 8.9 ± 2.4 1.3 ± 1.6 138.6 ± 37.3 7.4 ± 11.4 1.512 ± 0.197 0.411 ± 0.079
Nothofagus antarctica shrublands 12.2 ± 2.8 4.5 ± 2.5 113.4 ± 42.6 14.5 ± 19.7 1.716 ± 0.311 0.440 ± 0.130
Nothofagus antarctica krummholz 6.0 ± 2.1 - 136.6 ± 33.7 - 0.795 ± 0.371 0.728 ± 0.185
Pinus contorta plantations 3.3 ± 1.6 2.9 ± 0.7 26.6 ± 22.6 103.9 ± 41.9 0.879 ± 0.429 0.610 ± 0.239

149
Woodlands in the Chilean cold-temperate zone: VIDAL, O. ET AL.
TABLE II. Jaccard index values for all pairs of woodland communities.
TABLA II. Valores para el índice de Jaccard para todos los pares de comunidades comparados.
COMMUNITY TYPE 1 23456 7 891011
1. Nothofagus nitida forests 1.00 0.24 0.38 0.26 0.03 0.34 0.29 0.32 0.21 0.04 0.13
2. Pilgerodendron uviferum forests 1.00 0.37 0.15 0.05 0.14 0.22 0.18 0.10 0.07 0.07
3. Nothofagus betuloides forests 1.00 0.24 0.06 0.18 0.23 0.15 0.15 0.08 0.05
4. Nothofagus pumilio forests 1.00 0.06 0.45 0.35 0.29 0.43 0.05 0.14
5. Nothofagus pumilio krummholz 1.00 0.04 0.02 0.03 0.05 0.13 0.00
6. Nothofagus dombeyi forests 1.00 0.49 0.29 0.39 0.02 0.11
7. Embothrium coccineum shrublands 1.00 0.29 0.29 0.04 0.09
8. Nothofagus antarctica forests 1.00 0.25 0.05 0.26
9. Nothofagus antarctica shrublands 1.00 0.04 0.10
10. Nothofagus antarctica krummholz 1.00 0.04
11. Pinus contorta plantations 1.00
FIGURE 5. Map showing the location of phytosociological relevés in the study area: A) relevés belonging to the meso-hygromorphic
woodlands, B) relevés belonging to the hygromorphic woodlands and C) relevés belonging to the high-Andean dwarf woodlands. The
numbers indicates the towns close to the basins: 1) Caleta Tortel, 2) Cochrane and 3) the village of O’Higgins.
FIGURA 5. Mapa mostrando la locación de los relevamientos fitosociológicos en el área de estudio: A) relevamientos pertenecientes e las
comunidades leñosas meso-higromórficas, B) relevamientos correspondientes a las comunidades leñosas higromórficas y C) relevamientos
correspondientes a los matorrales achaparrados alto-andinos. Los números indican los poblados cercanos a las cuencas: 1) Caleta Tortel,
2) Cochrane y 3) Villa O’Higgins.

150
Gayana Bot. 68(2), 2011
A weak correlation between the number of exotics and
the distance to roads was observed (r= -0.213, p= 0.001)
when all relevés (n= 256) were evaluated, indicating
that roads have little influence as corridors for exotics.
Moreover, these correlations are stronger for some
communities when they are analyzed independently,
indicating specific responses of communities. Thus,
the presence of exotics in Nothofagus pumilio forests is
strongly correlated with the elevation of the relevés (r=
-0.494, p= 0.002) as well as with the distance to roads
(r= -0.493, p= 0.002). Similarly, Nothofagus antarctica
shrublands also showed significant correlations between
elevation of relevés and number of exotics (r= -0.454,
p= 0.001), but exotic presence in this community is not
significantly influenced by distance to roads.
DISCUSSION
COMPOSITION AND DISTRIBUTION OF WOODLAND COMMUNITIES
The native vegetation in the study area can be classified into
3 groups (Fig. 4): 1) Meso-hygromorphic woodlands, mostly
composed of deciduous woodlands with continental influence,
2) Hygromorphic woodlands, consisting of evergreen
woodlands with oceanic influence, and 3) High-Andean
dwarf woodlands, consisting of krummholz communities.
1) Meso-hygromorphic woodlands (Figure 5A), consisting
mainly of deciduous woodlands in warmer and drier places
in the NE section of the study area with sub-hyperoceanic
influence (Luebert & Pliscoff 2006), typically within the
Baker basin. The communities included here are N. antarctica
forests, E. coccineum tree shrublands, N. dombeyi forests,
N. pumilio forests and N. antarctica tree shrublands. The
vegetation in the Baker basin is more heterogeneous, consisting
mainly of a mosaic of vegetation formations with greater or
lesser degrees of disturbance due to livestock activities, but
its structure is less complex and contains a smaller number
of species as well as lower diversity. N. dombeyi forests were
described in the study area by Hambleton (1936), who located
these stands in the northern portion of the Baker basin. Other
communities from this belt, like N. pumilio forests and N.
antarctica forests, were also mentioned for this basin from
a phytogeographical point of view (Reiche 1934), but a
numerical comparison is hard to establish here. In this meso-
hygromorphic woodland group, N. dombeyi forests are the
only perennial community, which, as a typical component of
the northern-Valdivian phytogeographical region (Ramírez
et al. 1997, Amigo & Ramírez 1998), reaches its southern
distribution range in the study area. The southern limit of
this vegetation as a whole is the so-called ‘Cold Temperate
Zone’ (Godley 1960, Holdgate 1960, Skottsberg 1960), and
was established at 48°S by Godley (1960). Our southernmost
record for this belt reached 47.9°S.
2) Hygromorphic woodlands (Fig. 5B), consisting of
perennial woodlands in colder and more humid places in
the SW section of the study area, with euhyperoceanic
influence (Luebert & Pliscoff 2006), present at the southern
section of the Baker river basin and along the Pascua basin.
The communities included here are forests dominated by N.
nitida, forests dominated by Pilgerodendron uviferum and
forests dominated by N. betuloides. The vegetation is more
homogeneous than the meso-hygromorphic woodlands,
but its structure is complex, containing higher richness and
diversity than the drier belt (Amigo et al. 2004). Forests
dominated by N. betuloides were described by Hambleton
(1936), who extended their distribution to the archipelagic
areas, probably confusing the species with N. nitida. The
community dominated by N. nitida, however, was correctly
mentioned by Reiche (1934). It is situated in areas near
the mouth of the Baker River. N. nitida stands reach their
southern distribution limit at 49°S along the coastline, while
N. betuloides and P. uviferum stands reach the antiboreal
macrobioclimatic belt beyond 52°S (Oberdorfer 1960,
Amigo & Ramírez 1998, Luebert & Pliscoff 2006). Pisano
(1972), however, pointed out that it is not really possible
to speak of Pilgerodendron uviferum forests because of the
small size of the stands.
3) High-Andean dwarf woodlands (Fig. 5C), consisting
of deciduous krummholz communities at the snowline,
forming a continuous belt at upper elevations. Krummholz
dominated by N. pumilio are distributed in the Baker River
basin, while krummholz dominated by N. antarctica are
located in the Pascua River basin. Hambleton (1936)
mentioned the presence of krummholz communities in the
Baker basin above 1,000 m altitude, while Pisano (1972),
based on Davison’s data from New Zealand’s second
expedition to the North Patagonian Ice Field, described
N. antarctica krummholz communities with Empetrum
rubrum. This last species, however, has minor importance
with regard to frequency and cover in our plots.
DISTURBANCE PATTERNS AFFECTING WOODLAND COMMUNITIES
Four major types of current human disturbances affecting
the woodland communities have been observed: 1) logging
for fuel wood and construction timber, 2) overgrazing of
ground vegetation by livestock, 3) road widening in forested
lands and 4) invasion of some stands by Pinus contorta.
All of these disturbances are confined to the Baker basin
where roads, exotic plantations and human population
are present. The vegetation in the Pascua basin, however,
remains completely pristine. The disturbances occurring
in the Baker basin are slowly changing the structure and
composition of the woodland communities by means of
an increase in the introduction of exotic plant species,
reduction in cover of native dominant ones, mortality of
tree seedlings and erosion, among other effects, as occurs in

151
Woodlands in the Chilean cold-temperate zone: VIDAL, O. ET AL.
manuscript. Thanks to Juana Palma (Freiburg University)
for advising on the use of R software. The first two authors
thank the CONICYT and DAAD for the funding provided
to doctoral studies at the Albert-Ludwigs University in
Freiburg, Germany.
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1981, Raffaele et al. 2007, Quintanilla 2008, Speziale et al.
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species richness and diversity recorded for forests in the
Pascua basin.
Although our objective here is not directly to evaluate the
effects of past or current disturbances, it seems appropriate
to highlight the apparently low level of disturbances
affecting these forests, as we show through the biodiversity
indicators evaluated. This situation appears to offer even
an interesting opportunity for researchers to evaluate
undisturbed biogeographical and ecological patterns and
processes occurring at one of the most remote, isolated and
unpolluted landscapes in southern Chile.
ACKNOWLEDGMENTS
This project it was financed by the contract CA 012.05
UACH-UCONC-HIDROAYSEN. Thanks to Alejandra
Jiménez and Ernesto Teneb for field support. Prof. Dr.
Albert Reif (Freiburg University) and Prof. Dr. Jurgen Huss
(Freiburg University) commented on an earlier version of the

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ANNEX 1. Diagnostic species for each woodland community based on the indicator species values and p values (∗ ≤ 0.05; ∗∗ ≤ 0.005: ∗∗∗
≤ 0.001).
ANEXO 1. Especies diagnósticas para cada comunidad leñosa basado en los valores indicadores y sus correspondientes valores de probabilidad
p (∗ ≤ 0,05; ∗∗ ≤ 0,005: ∗∗∗ ≤ 0,001).
Community type / Species Family Origin Life Form ISA Value
Nothofagus nitida forests
Azara lanceolada Hook.f. Flacourtiaceae Native Phanerophyte 0.39 ***
Blechnum magellanicum (Desv.) Mett. Blechnaceae Native Hemicryptophyte 0.29 **
Campsidium valdivianum (Phil.) Skottsb. Bignoniaceae Native Epiphyte 0.41 ***
Drimys winteri J.R.Forst. et G. Forst. Winteraceae Native Phanerophyte 0.38 ***
Griselina ruscifolia (Clos) Ball. Griseliniaceae Native Epiphyte 0.56 ***
Hymenophyllum krauseanum Phil. Hymenophyllaceae Native Ephiphyte 0.70 ***
Lomatia ferruginea (Cav.) R. Br. Proteaceae Native Phanerophyte 0.23 ***
Mitraria coccinea Cav. Gesneriaceae Native Epiphyte 0.47 ***
Nothofagus nitida (Phil.) Krasser Nothofagaceae Native Phanerophyte 1.00 ***
Podocarpus nubigena Lindl. Podocarpaceae Native Phanerophyte 0.50 ***
Pseudopanax laetevirens (Gay) Franchet Araliaceae Native Phanerophyte 0.46 ***
Weinmannia trichosperma Cav. Cunoniaceae Native Phanerophyte 0.22 ***

153
Woodlands in the Chilean cold-temperate zone: VIDAL, O. ET AL.
Pilgerodendron uviferum forests
Berberis ilicifolia L.f. Berberidaceae Native Phanerophyte 0.40 ***
Carex magellanica Lam. Cyperaceae Native Hemicryptophyte 0.49 ***
Empetrum rubrum Vahl ex Willd. Empetraceae Native Chamaephyte 0.49 ***
Gaimardia australis Gaudich. Centrolepidaceae Native Chamaephyte 0.29 **
Hypochaeris palustris (Phil.) De Wild. Asteraceae Native Hemicryptophyte 0.29 **
Juncus microcephalus Kunth Juncaceae Native Hemicryptophyte 0.29 **
Lebetanthus myrsinites (Lam.) Dusen Epacridaceae Native Phanerophyte 0.48 ***
Macrachaenium gracile Hook.f. Asteraceae Native Hemicryptophyte 0.40 **
Myrteola nummularia (Poir.) O.Berg Myrtaceae Native Chamaephyte 0.31 **
Oreobolus obtusangulus Gaudich. Cyperaceae Native Hemicryptophyte 0.57 ***
Philesia magellanica J.F.Gmel. Philesiaceae Native Epiphyte 0.44 *
Pilgerodendron uviferum (D.Don) Florin Cupressaceae Native Phanerophyte 0.79 **
Nothofagus betuloides forests
Desfontainia fulgens D.Don Desfontainiaceae Native Phanerophyte 0.38 ***
Escallonia rubra (Ruiz et Pav.) Pers. Escalloniaceae Native Phanerophyte 0.24 ***
Hymenophyllum tortuosum Hook.et Grev. Hymenophyllaceae Native Epiphyte 0.22 *
Luzuriaga marginata (Banks et Sol. ex Gaertn.) Benth. Luzuriagaceae Native Epiphyte 0.30 **
Nothofagus betuloides (Mirbel) Oersted Nothofagaceae Native Phanerophyte 0.70 ***
Schoenus andinus (Phil.) H.Pfeiffer Cyperaceae Native Hemicryptophyte 0.20 *
Serpyllopsis caespitosa (Gaudich.) C.Chr. Hymenophyllaceae Native Epiphyte 0.50 ***
Sticherus quadripartitus (Poir.) Ching Gleicheniaceae Native Hemicryptophyte 0.22 **
Nothofagus pumilio forests
Adenocaulon chilense Poepp. ex Less. Asteraceae Native Hemicryptophyte 0.25 **
Codonorchis lessonii (Brongn.) Lindl. Orchidaceae Native Geophyte 0.18 *
Lycopodium magellanicum (P.Beauv.) Sw. Lycopodiaceae Native Hemicryptophyte 0.16 *
Maytenus disticha (Hook.f.) Urban Celastraceae Native Phanerophyte 0.43 ***
Nothofagus pumilio (Poepp. et Endl.) Krasser Nothofagaceae Native Phanerophyte 0.55 ***
Osmorhiza chilensis Hook et Arn. Apiaceae Native Hemicryptophyte 0.23 **
Poa pratensis L. Poaceae Exotic Hemicryptophyte 0.20 **
Viola maculata Cav. Violaceae Native Hemicryptophyte 0.49 ***
Nothofagus pumilio krummholz
Gaultheria pumila (L.f.) D.J.Middleton Ericaceae Native Chamaephyte 0.34 ***
Rubus geoides Sm. Rosaceae Native Hemicryptophyte 0.29 **
Nothofagus dombeyi forests
Nothofagus dombeyi (Mirb.) Oerst. Nothofagaceae Native Phanerophyte 0.89 ***
Ribes magellanicum Poir. Grossulariaceae Native Phanerophyte 0.18 *
Viola reichei Skottsb. Violaceae Native Hemicryptophyte 0.16 *
Embothrium coccineum shrublands
Baccharis magellanica (Lam.) Pers. Asteraceae Native Chamaephyte 0.49 ***
Embothrium coccineum J.R.Forst. et G.Forst. Proteaceae Native Phanerophyte 0.60 ***
Escallonia rosea Griseb. Escalloniaceae Native Phanerophyte 0.51 ***
Gaultheria mucronata (L.f.) Hook. et Arn. Ericaceae Native Phanerophyte 0.28 ***
Gavilea odoratissima Poepp. Orchidaceae Native Geophyte 0.34 ***
Lycopodium paniculatum Desv. Lycopodiaceae Native Hemicryptophyte 0.39 ***
Nothofagus antarctica forests
Blechnum penna-marina (Poir.) Kuhn Blechnaceae Native Hemicryptophyte 0.20 **
Chusquea culeou E. Desv. Poaceae Native Phanerophyte 0.76 ***
Escallonia virgata (Ruiz et Pav.) Pers. Escalloniaceae Native Phanerophyte 0.40 ***
Community type / Species Family Origin Life Form ISA Value

154
Gayana Bot. 68(2), 2011
Ranunculus trullifolius Hook.f. Ranunculaceae Native Hemicryptophyte 0.18 ***
Scirpus inundatus (R.Br.) Poir. Cyperaceae Native Hemicryptophyte 0.15 *
Nothofagus antarctica shrublands
Achillea millefolium L. Asteraceae Exotic Hemicryptophyte 0.26 *
Anemone multifida Poir. Ranunculaceae Native Hemicryptophyte 0.40 ***
Carduus nutans L. Asteraceae Exotic Therophyte 0.13 *
Cerastium arvense L. Caryophyllaceae Exotic Hemicryptophyte 0.16 *
Discaria chacaye (G.Don) Tortosa Rhamnaceae Native Phanerophyte 0.29 **
Festuca pyrogea Speg. Poaceae Native Hemicryptophyte 0.27 *
Fragaria chiloensis (L.) Duch. Rosaceae Native Hemicryptophyte 0.23 **
Geum magellanicum Pers. Rosaceae Native Hemicryptophyte 0.18 *
Jarava psylantha (Speg.) Penail. Poaceae Native Hemicriptophyte 0.16 *
Misodendrum punctulatum Banks ex DC. Misodendraceae Native Epiphyte 0.24 **
Phacelia secunda J.F.Gmel. Hydrophyllaceae Native Hemicryptophyte 0.17 *
Ribes cucullatum Hook. et Arn. Grossulariaceae Native Phanerophyte 0.18 **
Rumex acetosella L. Polygonaceae Exotic Hemicryptophyte 0.25 **
Nothofagus antarctica krummholz
Acaena pumila Vahl Rosaceae Native Hemicryptophyte 0.38 **
Bolax caespitosa Hombr. et Jacquinot Apiaceae Native Chamaephyte 0.60 **
Caltha dionaeifolia Hook.f. Ranunculaceae Native Hemicryptophyte 0.40 ***
Gunnera magellanica Lam. Gunneraceae Native Hemicryptophyte 0.62 ***
Marsippospermum grandiflorum (L.f.) Hook. Juncaceae Native Hemicryptophyte 0.30 **
Nothofagus antarctica (G.Forst.) Oerst. Nothofagaceae Native Phanerophyte 0.31 ***
Perezia linearis Less. Asteraceae Native Hemicryptophyte 0.18 *
Senecio acanthifolius Hombr. et Jacquinot Asteraceae Native Hemicryptophyte 0.92 ***
Senecio darwinii Hook et Arn. Asteraceae Native Chamaephyte 0.40 **
Pinus contorta plantations
Agrostis stolonifera L. Poaceae Exotic Hemicryptophyte 0.87 ***
Nertera granadensis (Mutis ex L.f.) Druce Rubiaceae Native Hemicryptophyte 0.51 ***
Pinus contorta Douglas ex Loudon Pinaceae Exotic Phanerophyte 1.00 ***
Community type / Species Family Origin Life Form ISA Value
Recibido: 28.02.11
Aceptado: 23.05.11