Tree Regeneration Strategies in a Lowland Nothofagus-Dominated Forest in South-Central

Chile
Author(s): Thomas T. Veblen, David H. Ashton and Federico M. Schlegel
Source: Journal of Biogeography, Vol. 6, No. 4 (Dec., 1979), pp. 329-340
Published by: Wiley
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Journalof Biogeography(1 97 9) 6, 3 29-340

Tree regenerationstrategiesin a lowlandNothofagus-

dominated forest in south-centralChile

THOMAST. VEBLEN, DAVID H. ASHTON*and FEDERICOM. SCHLEGEL

Departmento de Silvicultura,UniversidadAustral de Chile, Valdivia,Chile, and
*School of Botany, Universityof Melbourne,Australia

ABSTRACT.In the longitudinalvalley of south-centralChile remnantstandsof

old-growth forest on well-drained sites at altitudes of 100-400 m between
c. 390 and 410 S latitude are often characterized by scattered tall individuals of
Nothofagus obliqua and Eucryphia cordifolia which project approximately
10 m above a main canopy formed by Aextoxicon punctatum,Persealingue and
Laurelia sempervirens. The regeneration strategies of these tree species are
reflected by their size class and spatial distributionsand were studied in an old-
growth forest on the northwestern shores of Lake Villarrica.A. punctatum
regeneratesbeneath the shadeof the continuous forest canopy as well as beneath
small openings in the canopy. P. lingue and L. sempervirensregenerateonly
beneath small canopy openings, and E. cordifolia beneath both small and large
canopy openings. The regenerationof N. obliqua dependson complete or partial
destruction of old-growth stands. The regenerationstrategiesof the dominant
tree species form a continuumof responsesto differingscalesof disturbanceand
are appropriately perceived within a kinetic scheme of vegetation dynamics
emphasizingrepeateddisturbanceratherthan progressivedevelopmenttowardsa
stable end-point.

Introduction departures from the normal, but rather

Many studies of the structure and dynamics of
old-growth forests in the temperate zone have
demonstrated that individual stands are rarely
in a state of relative stability (e.g. Jones, 1945;
Sprugel, 1976). Nevertheless, classic
(Clements, 1916) as well as modern (Odum,
1969; Margalef, 1968) models of succession
describe succession as progressive change
towards an equilibrium community. In con-
trast to these 'developmental' schemes,
'kinetic' models of vegetation change do not
assume the existence of stable end-points to
succession (Drury & Nisbet, 1971). In a kinetic
scheme disturbances are not treated as unusual
Correspondence: Dr T. T. Veblen, Protection
Forestry Division, Forest Research Institute, P.O.
Box 31-011, Christchurch, New Zealand.
indeterminate change is accepted as an
essential part of complex, fluctuating systems.
Such a view is more consistent with the
repeatedly demonstrated instability of the
physical habitat (Raup, 1957, 1964) and is
adopted in this analysis, of the dynamics of a
lowland Nothofagus-dominated forest in
south-central Chile. Thus, the objective of this
study is the elucidation of the regeneration
strategies of the common tree species in
response to disturbances of differing scales
rather than the prediction of the species
composition and structure of the forest which
would theoretically develop in the absence of
disturbance. In the context of a closed canopy
forest, disturbance is defined as any phenome-
non resulting in an opening or, gap in the
canopy, whether it is the result of the gradual

0305-0270/79/1200-0329 $2.00 ? 1979 BlackwellScientificPublications 329

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330 T. T. Veblen, D. H. Ashton and F. M. Schlegel

death of a single large tree or group of trees, TABLE 1. Common species in the forest studied
or the physical levelling of a part of the forest.
Emergent tree stratum
Stand structure, or the frequency of Nothofagus obliqua (Mirb.) Bi.
differently sized individuals of tree species Eucryphia cordifolia Cav.
in a given stand, commonly reflects different Nothofagus dombeyi (Mirb.) Bi.
modes of tree reproduction. For example, an Matrix of dominant and subdominant trees
even-aged structure often suggests regenera- Aextoxicon punctatum R. et Pav.
tion following massive disturbances such as Persea lingue Nees.
catastrophic fire, whereas the populations of Laurelia sempervirens (R. et Pav.) Tul.
'stable' species capable of reproducing in their Small tree and shrub stratum
own shade should exhibit an all-age, or all-size, Gevuina avellana Mol.
Rhamnus diffusus Clos.
structure (Jones, 1945; Daubenmire, 1968). Lomatia dentata (R. et Pav.) R. Br.
Characterization of a species' population Myrceugenella apiculata (DC.) Kaus.
structure is usually based on the visual assess- Myrceugenia planipes (Hook. et Arn.) Berg.
ment of frequency distribution in age or size Amomyrtus luma (Mol.) Legr. et Kaus.
Lomatia ferruginea (Cav.) R. Br.
classes but may also be quantitatively assessed Caldcluvia paniculata (Cav.) D. Don.
(Hett & Loucks, 1976). The power function Rhaphithamnus spinosus (Juss.) Mold.
model has been demonstrated to describe the Peumus boldus Mol.
age or size structure of shade-tolerant species Chusquea quila (Mol.) Kunth
Aristotelia chilensis (Mol.) Stuntz
whose populations are in a steady-state condi- Fuchsia magellanica Lam.
tion in the sense that, in a given stand, there Lomatia hirsuta (Lam.) Diels
is a balance between the income of new
Lianas and epiphytes
individuals and the loss of individuals by Pseudopanax valdiviensis (Gay) Harms
death, thus maintaining a relative stability in Luzuriaga radicans R. et Pav.
total numbers (Hett & Loucks, 1976). The Hydrangea integerrima (Hook et. Arn.) Engl.
Hymenophyllum spp.
power function model is
Lapageria rosea R. et Pav.
Mitraria coccinea Cav.
y =yoXb
Boquila trifoliata DC.
where y is the number in any age or size Adiantum chilense Kaulf.
class x, yo is the initial input into the Cissus striata R. et Pav.
population at time zero, and b is the mor- Elytropus chilensis Muell.-Arg.
Lardizabala biternata Dcne.
tality rate. Polypodium feuillei Bertero
In interpreting regeneration strategies, the
Herbaceous ground stratum
spatial distribution of a tree species is often an Dysopsis glechomoides (Rich.) Muell.-Arg.
essential consideration and is particularly Rigodium implexum Kunz.
important in forests where a common mode Blechnum auriculatum Cav.
of regeneration is mass establishment beneath Osmorrhiza chilensis Hook. et Arn.
canopy gaps, which produces small clusters of Blechnum mochaenum Kunk.
Nertera granadensis (Mutis ex. L.f.) Drude
young trees (cf. Williamson, 1975). Conse- Blechnum chilense (Kaulf.) Mett.
quently, spatial as well as size class distribu- Lophosoria quadripinnata (Gmel.) C. Chr.
tions of the common trees are considered in Gunnera chilensis Lam.
this study of a lowland Nothofagus-dominated Oxalis dumetorum Gay
Hydrocotyle poeppigii DC.
forest in south-central Chile.

Study area derived from one to several metres of sandy

The forest studied is an old-growth forest
dominated in height by emergent individuals
of Nothofagus obliqua (see Table 1 for
botanical authorities) and is located on the
northwestern shores of Lake Villarrica (latitude
390 12'S, longitude 720 lO'W) at an elevation
of c. 250 m above sea level. The soil here is
volcanic ash and silty lacustrine deposits over-
lying glacio-fluvial materials. The 35 year
average annual rainfall, measured within a
kilometre of the forest studied, is 2900mm
with 27%, 45%, 20% and 8% falling during
autumn, winter, spring and summer, respec-
tively (Almeyda & Saez, 1958).

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 Regeneration strategies in a Nothofagus forest
40-42 m tall N. obliqua, the only
deciduous tree species present, project c. 10 m
above the main canopy level of the evergreen
tree species (Fig. la). The other common
emergent tree present is Eucryphia cordifolia
which attains heights of 35-40m; very few
emergent Nothofagus dombeyi (an average of
fewer than one in 5 ha) are also present.
Aextoxicon punctatum, Persea lingue and
Laurelia sempervirens form a dense matrix of
dominant and subdominant trees c. 30 m tall;
the most abundant tree, A. punctatum,
sometimes forms small pure stands c. 0.1 ha
in area (Fig. lb). Although rich in species
(Table 1), particularly in lianas, the under-
storey vegetation is very sparse; several
species such as the large climbing bamboo
Chusquea quila, the small shrub Fuchsia
magellanica, and the 4 m tall shrub Aristotelia
chilensis are found only along the edge of the
forest or beneath openings in the dense forest
canopy. The only evidence of direct human
intervention in this forest is a very few
scattered cut stumps, mostly of P. lingue; in
general, however, this forest has not been
exploited. The introduced red deer (Cervus
elaphus L.) has been present in this general
area for several decades and potentially could
seriously alter the structure of this forest by
destroying tree regeneration. However, the red
deer population is presently maintained at a
low level by hunting and its effects in the old-
growth forest are relatively slight due to the
availability of browse in adjacent secondary
vegetation and pastures.
On well-drained sites at altitudes of
approximately 100-400 m on the eastern
slopes of the coastal cordillera and on the
western slopes of the Andes between 390 and
410S in south-central Chile, the remnants of
forests floristically similar to that studied at
Lake Villarrica are common (Oberdorfer,
1960; Thomasson, 1963); the few known
stands which have not been severely altered
by fire or timber extraction are similar in
structure. Such forests probably occupied
most of the well-drained sites in the central
valley at these latitudes but have been cleared
to permit agricultural and pastoral land use;
today, only isolated trees, especially N.
obliqua and L. sempervirens, remain in
pastures.
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331
Methods
Vegetation sampling
For the analysis of stand structure, tree
diameters at breast height (dbh) of all
individuals > 5 cm dbh were measured in four
40 x 60 m plots which were subjectively
located to avoid trails and small stream
channels in stands where the main canopy
formed a relatively continuous cover. The
number of trees < 5 cm dbh but at least 2 m
tall (referred to as saplings in this study) were
also counted. Scales of tree clustering were
investigated by recording the dbh of trees
> 5 cm dbh and counting the number of
saplings in contiguous 5 x 5 m quadrats
forming a 10 x 3 20 m plot. This long plot
included mostly forest with a closed main
canopy but also passed beneath several canopy
openings caused by the recent death of indivi-
dual large trees; some of the gap-producing
large trees had fallen to the ground, typically
breaking off several metres above ground,
while others were still standing.
Five openings in the main canopy created
by the recent deaths of large trees (such as the
one depicted in Fig. 1c) were studied from the
point of view of their effects on tree repro-
duction. The numbers of saplings and of shrubs
beneath each canopy opening were recorded.
In each case the limits of the opening left in
the main canopy were mapped by vertically
projecting the crowns of all the surrounding
trees to the ground surface. The well-defined
patches of saplings and shrubs beneath each
canopy gap and extending slightly into the
adjacent undisturbed forest were also mapped;
the exact limits of the understorey vegetation
affected by the canopy gaps were easily deter-
mined because of the dramatically reduced
abundance of shrubs and saplings beneath the
adjacent continuous forest canopy. Numbers
of tree seedlings and shrubs < 2 m tall (but
excluding plants in the cotyledon stage) were
also determined in five random 1 x 2 m quad-
rats beneath each canopy opening and five
beneath the adjacent closed canopy.
Assessment of light conditions
Light conditions beneath openings in the
canopy and beneath areas of closed canopy
were assessed by the analysis of hemispherical
332 T. T. Veblen, D. H. Ash ton and F. M. Schlegel

photographs of the canopy of overstorey trees
(Anderson, 1964). One photograph was taken
at a height of 1.9 m from a centrally located
point beneath each canopy opening, and a
second photograph from a random point
beneath the adjacent closed canopy. The
percentage of total potential diffuse and
direct sunlight at a given site (i.e. compared
with total sunlight in the open over the same
period) was assumed to be approximately
equal to the percentage of visible sky; this
value corresponds to the 'total site factor' of
Anderson (1 964). The percentage of direct
potential sunlight, or the 'direct site factor' of
Anderson (1964), was estimated by
computing the percentage intercept of visible
sky along the sun's tracks for each month of
the year. Because the deciduous N. obliqua
occurs here as scattered emergent individuals
above the dense canopy of evergreen species,
it is assumed that seasonal changes in the
canopy density do not significantly affect
light levels measured at 1.9 m.
Data analysis
The size class distributions of tree species
were considered separately for the 10 x 3 20 m
plot and the four 40 x 60 m plots. Size
structure diagrams were prepared depicting

4m

Ls A Pi

(a)E

FIG. 1. Vegetation profiles of the forest studied: A, Aextoxicon punctatum; E, Eucryphia cordifolia;
Er, Eucryphia cordifolia root suckers; F, Fuchsia magellanica; G, Gevuina avellana; Lr, Lapageria rosea;
Ls, Laurelia sempervirens; Lu, Luzuriaga radicans; Mo, mosses; Mp, Myrceugenia planipes; Nd, Notho-
fagus dombeyi; No, Nothofagus obliqua; P1, Persea lingue; Pv, Pseudopanax valdiviensis; R, Rigodium
implexum; and Rs, Rhaphithamnus spinosus.

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 4m

(b) '

Mp MO

Rs~~~~~~~~~~

4m

A~~~~~~~~~~~~~~~

Pv~~~~

Nd

Er
Ls~~~~~R
L A F G~~~~~~~

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334 T. T. Veblen, D. H. Ashton and F. M. Schlegel
the frequency of saplings and of stems in
10 cm dbh classes beginning at 5 cm, The
power function model was used to evaluate
the departure from the ideal curve for a
steady-state population (Hett & Loucks, 1976)
where a size class distribution approached
the reverse J-shaped curve expected for a
continuously regenerating population
(Whittaker, 1974; Daubenmire, 1968). The
linear transformation of the power function
model used is
loge y = loge Yo - loge X
A least squares fit of this regression model to
the size class data was performed and the
departure from the expected ideal distribution
was assessed by the calculation of the correla-
tion coefficient.
For the analysis of spatial distributions of
the common tree species, mapping of all trees
in 5 x 5 m quadrats permitted application of
the nested quadrat technique described by
Greig-Smith (1964) and Kershaw (1973). The
measure selected for determining the departure
from a random distribution is Morisita's (1959)
index, the application of which has been
described by Williamson (1975). Morisita's
index, I6, is given by
q
Is = q L n (ni - lI)/N(N - l)
i=l
where q is the number of quadrats, ni is the
number of individuals of the species in the ith
quadrat and N is the total number of indivi-
duals in all q quadrats. The index, I8, equals
1.0 when the population is randomly distribu-
ted, where random implies an independent
distribution of individuals into quadrats with
an equal probability of each individual being
distributed in any one quadrat. If the indivi-
duals are clustered, Is is > 1.0, and if evenly
distributed or hyperdispersed, Is is < 1.0.
The possibility of determining the intensity
of clustering at a relatively large number of
small to intermediate size quadrats is the major
advantage of Morisita's index over the
commonly used analysis of variance technique
(Greig-Smith, 1964; Kershaw, 1973) in which
successive block sizes result from doubling the
size of the quadrat. Is was computed for
different species and different dbh classes
(< 10 cm and > 10 cm) for quadrats of
increasing size starting with quadrats of
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5x5m, 5x10m and lOxIOm, and subse-
quently increasing the 10 m wide quadrats by
10 m lengths up to a maximum quadrat size
of 10 x 80 m. Where the quadrat sizes are not
evenly divisible into the total area, I was
computed for the largest subareas containing
whole quadrats of a given size. In such cases,
the average index was computed from the
results obtained by initiating the analysis at
opposite ends of the 10 x 320 m plot.
Results and interpretation
Stand structure
Size class distributions of the five common
tree species are similar in both the combined
four 40 x 60 m plots (Fig. 2a) and the
l0 x 320 m plot (Fig. 2b). N. obliqua is repre-
sented exclusively by large individuals of at
least 50 cm dbh and clearly has not been
regenerating for some time in this old-
growth forest. On the other hand, the size-
class distribution of A. punctatum approaches
the ideal J-shaped curve expected for a stable
population; the power function model
describes its diameter distribution well in the
four 40 x 60 m plots (r = - 0.82;P < 0.01) and
in the 0x 320m plot (r=-0.87;P<0.01).
The other three species, P. lingue, E. cordifolia
and L. sempervirens, are represented by fairly
abundant saplings and small numbers of
individuals in all other size classes. Although
their size class distributions suggest all-age
populations, none approach the ideal distri-
bution described by the power function
model. Thus, the evidence of stand structure
reflects intermittent regeneration for P. lingue,
E. cordifolia and L. sempervirens, continuous
regeneration maintaining a steady-state popu-
lation of A. punctatum, and no recent
regeneration for N. obliqua.
Spatial facets
The number of shrubs and trees beneath
the five canopy gaps and the adjacent sites
beneath a closed canopy are shown in Tables
2 and 3. As expected, the numbers of tree
seedlings and shrubs are generally greater
beneath the canopy gaps with the exception
of P. lingue which is represented by approxi-
mately equal numbers of seedlings beneath
gaps and areas of continuous canopy. Although
 Regeneration strategies in a Nothofagus forest 335

2601

2504
200 7

190-

180-

170_

I20i' wo ( a) Aextoxicon punctatum

410

250
30-

20-

I0-

I 40
100- (ab) Aexrtoxi/con puJncOWUM

0 8718925-1 31933 loge

10ge y= X

sp I 2 3 4 6 7 8 9 1 II 12 13 14
20u u I
(r=--2 *
< 90 01 1! 2 1
2601U

1201H

least mtal,\an 1-14for tees blOcmb) cAextoxicon punctatum;

r 50
o Ioge Y=7?.7251-1.3933 o0ge X
40- (r=-0-87, P<OO01)
z
30-

20-

4 5 6 7
Sp 2 3 8 9 10 12
I II 13'
FIG. 2. Size structure diagrams and application of the power function model for the four 40 X 60 m
plots combined (a) and the 10 X 320 m plot (b). The size classes used are (sp) for trees < 5 cm dbh but at
least 2 m tall, and 1-14 for trees by 10 cm dbh classes beginning at 5 cm: n, , Aextoxicon punctatum;
L,Persealingue; El,Eucryphia cordifolia; ,Laurelia sempervirens; and U, Nothofagus obliqua.

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336 T. T. Veblen, D. H. Ashton and F. M. Schlegel

TABLE 2. Total numbers of shrub and tree seedlings distributions of the species' populations. Thus,
<2 m tall in 25 m2 samples beneath canopy openings throughout the forest initial establishment of
(>29m2 and <54 m2) and adjacent areas of con- some species beneath the canopy gaps should
tinuous canopy
result in relatively small clusters of younger
Species Beneath Beneath individuals. Whether clustering of the older
continuous canopy trees occurs or not depends on the amount
canopy openings of canopy space required by a mature indi-
Aextoxicon punctatum 123 890
vidual as well as on the size of the initial
Persea lingue 44 40 opening in the canopy. Even when the canopy
Rhamnus diffusus 10 20 gap is too small to permit the establishment
Eucryphia cordifolia 2 5 of more than a single large tree, clustered
Laurelia sempervirens 1 4
distributions of small individuals should occur
Lomatia dentata 1 2
Aristotelia chilensis 22 beneath the more recently created canopy
Nothofagus obliqua 6 openings. In Fig. 3 values of Morisita's index
Myrceugenella apiculata 3 are shown for trees > 2 m tall in size categories
Nothofagus dombeyi 1 of 6 10 cm and > 10 cm dbh; only populations
Lomatia hirsuta 1
Rhaphithamnus spinosus 1
with fourteen or more trees were analysed.
The dashed line in Fig. 3 at Is = 1.0 corre-
All species 181 995
sponds to a random distribution; filled symbols
are values of Is significantly greater than 1.0
(P < 0.05) according to an F-test of Morisita
TABLE 3. Mean number of saplings (>2 m tall and
<5 cm dbh) of tree species ha-' (? standard error) (1 959). Quadrat size is indicated in units of an
beneath canopy openings (>29m2 and < 54 m2) area of 5 x 5 m (i.e. as multiples of 25 mi2).
compared to sapling densities beneath the continuous The values of I6 given for P. lingue and E.
canopy cordifolia 6 10 cm dbh indicate highly
Species Beneath Beneath clustered distributions at small quadrat sizes
continuous canopy and a gradual tendency towards a random
canopy openings distribution at larger quadrat sizes; in other
words, small stems of these two species are
Aextoxicon punctatum 203?92 769? 303
56? 36 705? 343
contagiously distributed beneath the relatively
Persea lingue
Eucryphia cordifolia 15 ? 11 311 ? 174 small openings in the canopy. Although
Laurelia sempervirens 8? 7 330? 330 the L. sempervirens were not numerous enough
All species 282 ? 137 2115? 210 to permit the quantitative analysis of their
distribution, they also occur in clusters at
quadrat sizes 1 and 2. Sprouting from near the
L. sempervirens saplings were present in only base of the trunks of old individuals of L.
one of the five sites sampled beneath gaps, sempervirens and E. cordifolia as well as the
their presence was noted in several other vigorous root suckering of E. cordifolia also
similar sites not sampled. Abundant saplings contribute to the observed clustering of
beneath the canopy gaps and their relative saplings and small stems. However, successful
absence beneath the continuous forest canopy independent establishment of small indivi-
suggest that P. lingue, E. cordifolia and L. duals of these species following the death of
sempervirens regenerate only in canopy the parent tree was observed only beneath
openings. A few suppressed individuals of openings in the canopy. Poor dispersal of the
these species may persist beneath the con- heavy fruits (drupes) of P. lingue further
tinuous forest canopy where small canopy contributes to the observed contagious distri-
openings have recently closed up, but they bution of its saplings and small stems.
attain main canopy height only where a size- The values of I' for P. lingue > 10 cm dbh
able gap (i.e. 30-5Om2) persists in the main indicate strikingly even distributions at the
canopy. two smallest quadrat sizes and random distri-
A tree regeneration strategy in which butions at larger quadrat sizes. The change
young individuals establish principally beneath from a highly clustered distribution of small
canopy gaps should be reflected by the spatial stems to an even distribution of mature trees

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 Regeneration strategies in a Nothofagus forest 337

2-4 at the two smallest quadrat sizes suggests that,

of the numerous saplings of P. lingue beneath
22 r a small canopy opening (c. 25-50 m2), only a
(a ) Aextoxlcon punctotum
2 0/ single individual survives to maturity. E.
cordifolia > 10 cm dbh were not numerous
I.e . enough to permit analysis of their distribution
1 6- but are not noticeably clustered.
The values of Is for A. punctatum < 10 cm
14 dbh indicate a contagious distribution for all
12 quadrat sizes analysed with maximal clustering
at the largest and smallest quadrat sizes,
I0* although the intensity of pattern is relatively
low. The peak at quadrat size 32 reflects the
O. I . . . noticeably greater abundance of 5-10 cm dbh
stems in one-third of the 10 x 320 m plot
where an extensive area of the forest appeared
to have been more recently affected by a
7 0-
massive disturbance than the remainder of the
6*0 - (b) Eucryphla cordolfo/la forest. The small peak at quadrat size 1 for
small stems of A. punctatum may reflect not
5- 0- only a response to small openings but also
5*0 vigorous vegetative reproduction (both by
basal sprouting and root suckering) and
3 -0- the poor dispersal of its heavy fruits (drupes).
240-
*0-
In general, however,A. punctatum 6 10 cm dbh
\O
are much less clustered at the smaller quadrat
sizes than are E. cordifolia and P. lingue, which
suggests that this species is less dependent on
small openings in the canopy for successful
establishment. A. punctatum > 10 cm dbh are
8 0- randomly distributed at all quadrat sizes.
(c) Persea lingue

6 - 0- Light conditions
5 *0- Mean percentage total sunlight (diffuse and
direct) beneath the canopy gaps is 45.9 ? 4.3 SE
compared to 11.9?0.7 SE beneath the con-
tinuous forest canopy. During late spring and
2-0- early summer (November-January) mean per-
centage direct sunlight is much greater
beneath the canopy gaps than beneath the
0 / continuous forest canopy (Fig. 4); however,
4 8 12 16 20 24 28 32 the differences in direct sunlight beneath gaps
Quadrot size and the continuous canopy during winter and
autumn are very minor because of the low
FIG. 3. Values of Morisita's index, I,j, at different angle of the sun's path.
quadrat sizes for (a) Aextoxicon punctatum ?10 cm Least-squares linear and logarithmic regres-
(-, o) and A, punctatum >10cm (-, Q), (b) sions fail to demonstrate a significant relation-
Eucryphia cordifolial10cm (-, 0), and (c) Persea ship (P < 0.0 5) between the percentage total
lingue610cm (0, 0) and P. lingue>lOcm (-, 0).
Filled symbols are I,6 values significantly greater than sunlight beneath each canopy gap and the
1.0 (P < 0.05) according to an F-test of Morisita number per ha of tree saplings of all species
(1959). Quadrat size is indicated in units of 25 m2. combined. Similarly, tree sapling density and

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338 T. T. Veblen, D. H. Ashton and F. M. Schlegel

-
50-

'40-

. 50

( 10 0 < ~~0

2: 0-
J A S 0 N D J F M A M J
Months of the yeor

FIG. 4. Monthly variation in mean percentage direct sunlight for five sites beneath canopy openings
>29m2 and <54m2 (0, 0) and five adjacent sites beneath continuous canopy (M, 0). Filled symbols
are values significantly different (P < 0.01; t test) for the two types of sites being compared.

the area of each canopy gap are not signifi-
cantly correlated. On the other hand, tree
sapling density is significantly correlated
with mean monthly percentage direct sunlight
(r = 0.79; P < 0.01). If percentage direct
sunlight of only the months of the vegetative
period (October-April) are considered, the
strength of this relationship is greater (r=0.88;
P < 0.001), suggesting that successful estab-
lishment of tree saplings depends less on the
total size of a canopy gap than on its confi-
guration and orientation which control the
amount of direct sunlight reaching the under-
storey during the vegetative period.
Regeneration strategies and reproductive
characteristics
Stand structure and spatial distributions of
the common tree species in this N. obliqua-
dominated forest reflect three principal strate-
gies of tree regeneration. E. cordifolia, P. lingue
and L. sempervirens regenerate beneath small
canopy openings such as those created by the
death of a single large tree; under the much
lower light levels beneath the continuous forest
canopy they generally fail to achieve sapling
size. Even though A. punctatum can regenerate
beneath the shade of the continuous forest
canopy, saplings and small stems are more
abundant beneath canopy gaps than where
the canopy is continuous, thus indicating the
favourable effect of small canopy openings on
its regeneration. N. obliqua completely fails
to regenerate both beneath small openings
in the canopy as well as beneath the closed
canopy; its regeneration requires extensive
open areas created by massive disturbance. In
the surrounding area where the forest has
been burned or clear-cut N. obliqua charac-
teristically is the most abundant and domi-
nant tree in the secondary vegetation. E.
cordifolia is also an important component of
stands which have recently originated from
the burning or clear-cutting of old-growth
forests, whereas the relative abundances of
P. lingue, L. sempervirens and A. punctatum
are greatly reduced in young secondary
forests.
The strategies of tree regeneration inferred
from the structure of the Lake Villarrica
forest basically reflect the differences in the
common tree species' capacities to grow under
the environmental conditions associated with
understorey position in a dense forest. The
understorey tolerance (sensu Spurr & Barnes,
1973) of A. punctatum is the greatest and
that of N. obliqua the least while those of
P. lingue, L. sempervirens and E. cordifolia are
intermediate. In addition to understorey
tolerance, a tree's capacity to disperse its
disseminules and to reproduce vegetatively
are important aspects of its overall regenera-
tion strategy, and in the case of the common
tree species in this forest these reproductive
traits are essentially consistent with the
regeneration strategies previously outlined.
Dissemination of E. cordifolia and N. obliqua
to open sites is assured by their small, winged
seeds; on primary bare areas, such as landslide
surfaces in the foothills of the Andes, they
establish in large numbers and grow relatively
rapidly (Veblen & Ashton, 1978). Where the
old-growth forests have been heavily exploited,
both N. obliqua and E. cordifolia sprout

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 Regeneration strategies in a Nothofagus forest
vigorously from cut stumps in addition to
reproducing abundantly from seed. Although
seedlings of E. cordifolia are scarce beneath
a continuous forest canopy (Table 2) probably
because of their relative intolerance to shading,
basal sprouting and root suckering by this
species permit occasional regeneration beneath
canopy gaps in the old-growth forest at Lake
Villarrica. No seedlings of N. obliqua establish
beneath a closed forest canopy although a few
are found beneath canopy openings; under
canopy openings produced by the death of
one large tree or a small group of trees, this
shade-intolerant species does not attain sapling
size. The regeneration of L. sempervirens
beneath canopy gaps is favoured by the
capacity of old individuals to produce basal
sprouts. Furthermore, its light comose seeds
are efficiently dispersed by wind, thus
increasing the chances of the seed arriving at
sites beneath canopy gaps. Where observed in
forests of similar composition in the coastal
cordillera of south-central Chile, the seedlings
of L. sempervirens appear to be relatively
shade-tolerant and their scarcity in this old-
growth forest is puzzling. The ground layer of
moss and litter may impede contact between
its light seeds and the mineral soil. On the
other hand, both P. lingue and A. puncta tum
produce heavy disseminules and are represen-
ted by abundant small seedlings beneath the
continuous forest canopy. These heavy
disseminules are dispersed short distances by
gravity and longer distances by birds. Seedlings
of P. lingue achieve sapling and tree size
principally, if not exclusively, beneath canopy
gaps created by the death of one or more trees
in the main canopy. Its regeneration strategy
differs from that of the other 'gap species' in
that it generally does not reproduce vegeta-
tively. In contrast, A. punctatum vigorously
reproduces by means of basal sprouting and
root suckering as well as from seed. Although
its regeneration is enhanced beneath small
canopy gaps, it also regenerates beneath a
relatively continuous forest canopy.
Conclusions
The dynamics of the lowland N. obliqua-
dominated forest studied at Lake Villarrica
are analogous to the dynamics of Nothofagus-
dominated forests of the mid-montane zone
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339
of the Andes of south-central Chile. Neither
of the two emergent species which dominate
the mid-elevation forests, N. dombeyi and N.
alpina (Poepp. et Endl.) Oerst., regenerate
beneath a closed forest canopy nor beneath
small openings in the canopy; the regeneration
of both species depends on massive distur-
bances such as the periodic catastrophic mass
movements and vulcanism typical of this
region (Veblen & Ashton, 1978). Similarly, in
the near-timberline zone of the Andes, the
structure and regeneration of the pure N.
pumilio (Poepp. et Endl.) Krasser forests and
mixed forests of this species with N. dombeyi
and N. betuloides (Mirb.) Bi. are also greatly
influenced by mass movements and vulcanism
as well as by snow avalanches and windstorms
(Veblen et al., 1977). The structure of the
lowland forest at Lake Villarrica suggests
massive establishment of N. obliqua at least
250 years ago under open conditions created
by some type of extensive disturbance. Prior
to European colonization in the mid-nine-
teenth century, this area was occupied by an
aboriginal population (as indicated by native
graves beneath the present forest) which prac-
tised a rudimentary agriculture that required
burning of the forest (Lauer, 1961; Encina,
1954). Although direct evidence is not
available, burning of the vegetation by the
native population could account for the
present forest structure in which the shade-
intolerant N. obliqua dominates a matrix of
several more shade-tolerant tree species. Fire
set by the aboriginal population is the most
probable explanation for the structure of the
forest studied at Lake Villarrica, but wide-
spread forests of similar composition and
structure at low elevations in south-central
Chile may also be attributed to disturbances
such as severe windstorms, lightning-initiated
fires, volcanic ash deposition, catastrophic
flood damage, and, where relief is greater,
mass movements. Although these catastrophic
phenomena are infrequent in terms of a
human life span, they are relatively frequent
during the several hundred years required for
the development of these old-growth forests.
In a developmental scheme of plant succes-
sion the forests studied at Lake Villarrica
would be interpreted as a seral stage in the
slow, progressive development in a stable
physical habitat towards a relatively stable
340 T. T. Veblen, D. H. Ashton and F. M. Schlegel
forest dominated by the more shade-tolerant
species. However, it is more likely that this
forest would be affected by minor and massive
disturbances before such a relatively stable
forest develops. Thus, it seems more useful to
view this forest within a kinetic scheme of
succession in which the present structure and
species composition are regarded as conse-
quences of the differential responses of the
dominant species to periodic disturbances of
varying scales.
Acknowledgments
For preserving the forest studied and for their
cordial hospitality we are grateful to the
Ernesto Wagner family. We thank A. T. Veblen
for drawing the Figures and providing helpful
comments on the manuscript.
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