Journal of Vegetation Science 10: 861-868, 1999

IAVS; Opulus Press Uppsala. Printed in Sweden

- Tree regeneration and future dynamics of the laurel forest on Tenerife -

861

Tree regeneration and future dynamics of the laurel forest

on Tenerife, Canary Islands
Arvalo, Jos Ramn1*, Fernndez-Palacios, Jos Mara1 & Palmer , Michael W.2
1Departamento

de Ecologa, Universidad de La Laguna, La Laguna 38206, Tenerife, Spain;

of Botany, Oklahoma State University. Stillwater, OK 74078, USA;

2Department

*Corresponding

author; Fax +34922318311; E-mail: jarevalo@ull.es

Abstract. We studied two sites in the laurel forest of Tenerife

to predict future changes in canopy composition. We used two
projection methods: Horns Markovian Projection, which
utilizes information on juveniles in the vicinity of canopy
trees, and a Stand Projection, which ignores such information. We performed these projections both including and excluding a-sexual regeneration. Although all of our projections
predict a change in species composition, inclusion of a-sexual
reproduction decreased the magnitude of successional change.
The persistence of Prunus lusitanica and Ilex canariensis
appears to be highly dependent on a-sexual regeneration.
Both the Markov- and stand projections predict a slight
convergence in species composition between the two sites
when only sexual regeneration is considered, and also a higher
dominance of the shade-tolerant species Laurus azorica. When
a-sexual regeneration is also considered, some divergence is
shown, with less projected change in the stand projection than
in the Markov projection. In spite of some differences between
the models, general patterns such as an increase of shadetolerant species (Laurus azorica and Prunus lusitanica) and a
decrease of shade-intolerant species (Erica arborea, Erica
scoparia and Myrica faya) are consistent.

The successional status of the laurel forests of

Macaronesia is poorly known. While it appears that
differences among tree species with respect to shade
tolerance, mode of reproduction, seed bank, and spatial
pattern are likely to drive the dynamics of laurel forest,
long-term composition trends have not been documented
(Arvalo 1998; Arvalo & Fernndez-Palacios 1998).
While the best understanding of forest dynamics comes
from long-term observation (e.g. Busing 1996), model
predictions of future composition can be used to investigate the likelihood of change from present observations.
In this paper, we explore the likely dynamics of
laurel forest. We performed a separate analysis with and
without sexual and a-sexual reproduction to assess the
consequences of reproductive mode for future composition. We asked the following questions: is the laurel
forest likely to change through time and, if so, is there any
evidence for successional convergence across sites? Are
predictions sensitive to the model employed? Are predictions based on sexual and a-sexual reproduction different
than predictions based on just sexual reproduction?

Keywords: Basal sprout; Canopy dynamics; CCA; DCA;

Markov projection; Sapling; Stand projection.

Material and Methods

Nomenclature: Hansen & Sunding (1985).

Study site
Introduction
Models that predict directional changes in plant species composition are useful tools that can help us understand the directions, rates, and mechanisms of such
changes (e.g. Gillman & Hails 1997). Two of the most
commonly used models are Markov projection (Horn
1975, 1981) and stand projection (Hubbell & Foster
1987). These two models use different approaches and
predict composition at different points in the future.
Markov models use understorey densities beneath each
species of canopy tree and project composition at equilibrium. Stand projection models use summed
understorey density for an entire stand and predict composition in the next generation of canopy trees.

The laurel forest of Tenerife has been extensively

exploited since the arrival of Europeans in the 15th century
(Parsons 1981). The remaining forest (ca. 10 % of the
original area) is relatively well conserved and has experienced only minimal human disturbance in the last 50 yr.
The laurel forest was formally protected in 1988. No data
are available about the age of the forest but we assume that
it is at least a few generations old. Although illegal human
disturbance processes occurred until 50 yr ago creating
some earlier successional patches in the forest, these were
minor and did not affect the study sites. This assumption is
supported by the lack of government exploitation reports
from this area for the last three centuries. Aerial photographs from 1952 show a similar forest (in extent and
physiognomy) without noticeable disturbance.

862

Arvalo, J.R. et al.

Data collection

More than 95 % of the regeneration of some tree

species in the laurel forest is a-sexual (primarily by basal
sprouts). The resulting dominance by a few species is
characteristic of remnant communities (Eriksson 1996)
and a low disturbance level (Packham et al. 1992). The
seed bank of the forest is constantly replenished through
the seed rain, and is dominated by shade-intolerant species with a long residence time (Arvalo 1998). The role
of the seed bank in the succession is most likely determined by the disturbance regime. However, in previous
studies we found no significant differences in species
composition of regeneration between gap and understorey
(Arvalo & Fernndez-Palacios 1998). Furthermore, we
found no significant differences in the regeneration composition among different sites with different canopy composition (Fernndez-Palacios & Arvalo 1998).
The study was located in the Anaga Natural Park,
NE Tenerife (28o 19' N, 16 o 34' W). The park encompasses a 7 to 8 million-yr-old basaltic massif (Ancochea et
al. 1990) covering ca. 130 km2. The park represents 7 %
of Tenerifes area. We selected two stations in the park
as representing the best-conserved laurel forests of
Anaga: El Moquinal on the windward slope and Monte
de Aguirre on the leeward slope. We chose ca. 300 ha
of the most intact forest in each station (Table 1).
The annual precipitation of the park reaches 900
mm, but can be twice this amount if fog drip is included
(Kmmer 1974). Mean annual temperature is ca. 15 C
with minimal annual and daily fluctuations. There is no
frost. Soils have been classified in the order Entisol,
suborder Orthens (Fernndez-Caldas et al. 1985).
The canopy of Anagas laurel forest varies between
10 and 20 m high depending on slope. Maximal heights
are found at the basins floor decreasing progressively
towards the basins borders. Dominant species include
Laurus azorica, Erica scoparia, E. arborea, Ilex canariensis, Prunus lusitanica and Myrica faya. Viburnum
tinus is an understorey tree which rarely reaches the
canopy; it is not considered a canopy tree in this study.
The laurel forest contains a total of 19 tree species
(Santos 1990). Further information about the study site
is given in Fernndez-Palacios et al. (1992) and
Fernndez-Palacios & de Nicols (1995).

In June and July of 1996, we randomly selected four

new 625-m2 square plots in each of the two sites with
different altitude and aspect (plots 1 to 4 were in Moquinal
and plots 5 to 8 in Aguirre). Because we established new
plots, we should be cautious with the extrapolations of
results from our earlier studies (Arvalo & FernndezPalacios 1998; Fernndez-Palacios & Arvalo 1998).
At each plot we noted slope, aspect, altitude, UTM
(Universal Transverse Mercator grid) coordinates, percentage understorey cover and canopy cover (Table 1).
We defined trees as stems of at least 4 cm DBH (diameter at breast height), saplings as stems taller than 50 cm
and less than 4 cm DBH and originating from seeds, and
basal sprouts as sapling-size stems of a-sexual origin
(connections with parent stems, with some exceptions,
were apparent). We mapped all trees, saplings and basal
sprouts to an accuracy of 0.05 m.
Analyses of predictions
Two methods have been extensively used for predicting future canopy composition: Markov projection
and stand projection. The Markov projection assumes
that the proportion of saplings belonging to a particular
species beneath a canopy tree is an estimate of the
probability of replacement by that species (Horn 1975;
van Hulst 1979; Orlci & Orlci 1988). Some objections have been raised against this model, especially due
to the assumption of a constant transition matrix (Lippe
et al. 1985), a problem that could be minor when the
forests are close to the steady state (Horn 1975). When it
is not possible to obtain historical information about the
composition of the forest, low order Markov models can
reveal the direction of change in species composition
(Dale et al. 1993).
For the Markovian projection, we subdivided the
plot into Vornoi polygons (each polygon represents the
region of the plots closer to the tree of interest than to
other trees), one for each tree, and counted the number
of saplings in each polygon. We summed the sapling
counts over all polygons for each canopy species. We

Table 1. General biotic and abiotic information on the plots. Plots 1 to 4 are from the Moquinal site and 5 to 8 from the Aguirre site.
Abiotic characteristics
Altitude
Exposition
Slope(o)
(m)
Plot 1
2
3
4
5
7
8

770
705
785
755
965
945
910

N
NW
NW
NW
S
S
S

20
35
20
17
30
30
30

Canopy
Cover (%)

Understory
Cover (%)

100
95
100
100
95
95
95

15
5
10
15
5
5
15

Biotic characteristics
Basal area
Saplings
(m2/ha)
(stems/m2)
34.40
43.16
37.04
40.93
54.04
46.82
44.29

0.10
0.02
0.04
0.10
0.19
0.08
0.16

Suckers
(stems/m2)
0.25
0.15
0.36
0.49
0.13
0.14
0.36

- Tree regeneration and future dynamics of the laurel forest on Tenerife then constructed a transition probability matrix from the
proportion of saplings under each canopy species. The
transition matrix is multiplied by a vector that represents
the proportions of species densities in the plot. The
product is a second vector that represents the proportions of species densities in a theoretical time +1
(which loosely corresponds with the time a tree remains
in the canopy), which we multiply again with the transition probability matrix. After many iterations we reach a
stationary vector, that represents the proportions of species densities in the steady state.
We performed an additional Markov analysis in
which the transition probability matrix was calculated
from saplings plus basal sprouts. Because the transition
matrices differ, we obtained two different Markov projections per plot, one from sexual regeneration and
another from sexual plus a-sexual regeneration.
The stand projection method assumes that the proportion of saplings in the entire stand will estimate the
proportion in the future canopy (Hubbell & Foster 1987).
This projection does not predict a steady state, only
changes in the next generation of the forest. Also, this
model does not assume that dynamics are influenced by
the spatial distribution of individuals within the stand.
Because the two models differ in their assumptions, we
expect that they will differ in their predictions, depending on the importance of site effects and the spatial
relationships of canopy replacement. As with the
Markovian projection, two different projections were
obtained from this method, one from sexual regeneration
and another one from sexual plus a-sexual regeneration.
Both methods assume that the probability of a sapling
reaching the canopy is independent of the type of species.
Another assumption is that species have similar life spans.
We do not have data on longevity in the laurel forest, but
similar studies correcting for differences in longevity did
not reveal fundamentally different results (Horn 1975).
However, other studies reported the critical role of longevity in the dynamics of the forest (White et al. 1985).
Preliminary studies in the laurel forest (Arvalo
1998) revealed three canopy species to be shade-intolerant or pioneers (Erica arborea, E. scoparia and Myrica
faya), two species to be shade-tolerant or secondary
species (Laurus azorica and Prunus lusitanica), and one
species to have an intermediate tolerance (Ilex canariensis).
These six species represented ca. 95 % of the total canopy
composition and ca. 90 % of the total regeneration (Table
2). Less frequently occurring species (Apollonias barbujana, Ilex platyphylla, Persea indica, Picconia excelsa,
Rhamnus glandulosa and Teline canariensis) were assigned to a category Other species.
We used the paired Student t-test to compare the
mean of the percentage of densities per species with the
projected mean for each station (four present plots vs.

863

four projected plots). The use of the proportions created

a numerical dependence between species. Also, there is
a built-in dependence between predicted and observed
species composition, since the latter is derived from the
former. These unavoidable problems limit the validity
of statistical inference, so we view such inference to be
provisional and exploratory.
Multivariate analysis
We used Detrended Correspondence Analysis (DCA,
Hill & Gauch 1980) to analyse present and projected
species composition. Ordination is not only useful in
assessing the amount of change, but also directionality
and the nature of change. For example, if plots moved in
different directions in ordination space, this would imply
successional divergence.
We performed four Canonical Correspondence
Analyses (CCA; ter Braak 1986) based on the results of
the four different projections. There was only one explanatory variable: a dummy variable that represents
whether the sample was observed or projected. Because
there is only one variable, there is only one CCA-axis.
Species scores along this axis represented the degree to
which the species is projected to increase in the future
(i.e. represented in the projected samples). Therefore,
the axis is a convenient index of inferred successional
rank of species under different models. We also performed CCA with the stations (Moquinal vs. Aguirre) as
the only explanatory variable. We tested the existence of
differences between stations and whether such differences were maintained in the four different projections, or
whether the succession led to convergence across sites.
We performed all multivariate analyses with the
CANOCO package (ter Braak & milauer 1998) and
tested the eigenvalue of the axis with a Monte Carlo Test
using 200 iterations.
Except where otherwise noted, statistical methods
followed Zar (1984) and were implemented using SPSS
(Anon. 1986).

Results
Tree species composition
The canopy of the plots of Moquinal was, with
respect to basal area and density, dominated by Prunus
lusitanica and Ilex canariensis, and in Aguirre by Myrica
faya, Erica scoparia and Laurus azorica (Table 2).
Saplings of L. azorica occurred in all plots. Basal sprouts
of P. lusitanica and L. azorica dominated in Moquinal
and basal sprouts of L. azorica dominated in Aguirre.
Although P. lusitanica is a species with a high rate of

864

Arvalo, J.R. et al.

Table 2. a. Basal area/ha; b. Density/ha; c. No. of saplings/100m2 and d. No. of suckers/100m2 by species and plots. Other species
are: Apollonias barbujana, Ilex platyphylla, Persea indica, Picconia excelsa, Rhamnus glandulosa and Teline canariensis.
a. Basal area
Erica arborea
Erica scoparia
Ilex canariensis
Laurus azorica
Myrica faya
Prunus lusitanica
Other species

Moquinal site plots

1
8.76
0.66
5.26
3.67
4.04
8.19
3.82

2
4.49
9.98
10.65
17.98
-

b. Density

4
2.19
8.75
4.95
6.71
14.70
4.16

Mean
2.64
0.75
7.11
5.77
6.44
13.89
2.31

Std
3.61
0.87
2.29
2.48
2.64
3.55
1.74

5
0.12
23.13
2.94
6.60
20.65
0.67

6
0.29
13.84
5.48
8.53
16.85
0.01
0.82

Moquinal site plots

1
Erica arborea
256
Erica scoparia
Ilex canariensis 1200
Laurus azorica
688
Myrica faya
144
Prunus lusitanica 1264
Other species
352

2
432
544
208
960
-

c. No. of saplings / 100m2

Erica arborea
Erica scoparia
Ilex canariensis
Laurus azorica
Myrica faya
Prunus lusitanica
Other species

3
1.78
0.15
9.93
4.46
4.36
14.68
1.25

Aguirre site plots

3
32
16
1232
720
80
2256
64

4
64
912
544
160
1296
192

6.08
0.16
0.80
3.20

1.76
-

0.32
3.36
0.64
-

0.32
7.68
0.48
1.12

d. No. of suckers /100m2

Mean
0.16
4.72
0.04
0.48
1.08

Std
107.0
26.2
321.8
81.8
46.8
487.0
135.6

5
16
3728
400
1136
1216
96

6
48
1568
528
704
512
48

2.08
4.32
1.44
7.36
-

3
4.32
5.12
26.88
0.32

4
8.96
10.08
0.96
29.12
1.44

Mean
4.36
7.68
0.60
18.84
0.76

Mean
3.16
12.59
4.44
9.08
16.46
0.02
1.51

Std
4.94
8.22
0.96
2.27
3.06
0.02
0.83

7
32
1088
480
864
400
192

8
528
16
384
1264
256
16
112

Mean
156.0
1600.0
448.0
992.0
596
4.0
112.0

Std
215.1
1351.0
59.8
220.3
369.3
7.9
52.8

Aguirre site plots

Std

0.16
2.30
0.07
0.30
1.31

0.32
0.16
14.72
0.48
0.32
3.20

6
17.60
0.16
3.04

Moquinal site plots

2

8
11.71
0.05
4.31
12.78
12.03
0.05
1.81

Aguirre site plots

Moquinal site plots

1
Erica arborea
Erica scoparia
Ilex canariensis
2.08
Laurus azorica 11.20
Myrica faya
Prunus lusitanica 12.00
Other species
1.28

Mean
72.0
20.0
944.0
624.0
148.0
1444.0
152.0

7
0.51
13.35
5.01
8.40
16.32
2.73

7
0.32
0.32
5.12
0.16
0.32
1.28

8
11.36
0.64
2.88

Mean
0.16
0.12
12.20
0.16
0.36
2.60

Std

Mean
1.00
2.40
11.68
2.92
0.12
0.36

Std

0.16
0.13
4.65
0.20
0.17
0.77

Aguirre site plots

Std

2.81
3.00
0.62
9.34
0.61

2.56
2.88
9.28
3.68
-

0.64
2.56
4.80
3.68
-

0.80
2.88
9.60
1.28
0.80

regeneration by basal sprouts which is related to the

ecological strategy of regeneration (Arvalo & FernndezPalacios 1998) its low density in Aguirre site is the
origin of the differences of a-sexual regeneration among
sites (Table 2).
Projection of future canopy composition
The two projection methods had similar predictions
when considering only sexual regeneration (Fig. 1a, c).
Ilex canariensis and Prunus lusitanica were projected to
decrease while Laurus azorica was projected to increase
in density in Moquinal. I. canariensis and Myrica faya
are projected to decrease while L. azorica and Other
species were projected to increase at Aguirre. Results
differed between sites when a-sexual reproduction was
included in the projection (Fig. 1b, d). At Moquinal the
Markov projection indicated a significant decrease in I.
canariensis and an increase in L. azorica while the stand
projection only revealed a decrease in I. canariensis. In

8
1.28
23.04
3.04
0.48
0.64

0.95
0.66
6.83
0.98
0.21
0.36

Aguirre, the Markov projection did not produce any

significant differences between present and projected
densities of species while the stand projection indicated
an increase of L. azorica and Other species.
DCA reveals the successional trajectories projected
by various methods (Fig. 2). We present separately the
species scores (Fig. 2a) while we present the scores of
the plots of the four different projections together with
the scores of the present plots. An arrow represents the
projected trajectory of the plots through succession. The
trajectories by both methods using just sexual regeneration are similar (Fig. 2b, d). L. azorica and Other species
increased in importance. Shade-intolerant species Erica
arborea, E. scoparia and M. faya decreased, along with
Prunus lusitanica (shade-tolerant) and I. canariensis (intermediate behaviour). The projections by both methods
including a-sexual regeneration were more heterogeneous, but both indicated a decrease in shade-intolerant
species. When a-sexual regeneration is incorporated in
the projections, P. lusitanica and I. canariensis showed

- Tree regeneration and future dynamics of the laurel forest on Tenerife AGUIRRE
Sapling based projections

MOQUINAL
Sapling based projections
*

Pl

Mf

Species

Present

Present

Markov projection

Markov projection

Stand projection

Stand projection

*
*

La

Ic

865

* *

Es

Ea

*
*

OS

20

40

60

80

100

MOQUINAL
Sapling and sucker based projections

AGUIRRE
Sapling and sucker based projections

Present
Pl

Present
Pl

Markov projection

Markov projection

Stand projection

Stand projection

Mf

Mf

Ic

La

*
Species

Species

La

Ic

Es

Es

Ea

Ea

OS

OS

*
0

20

40

60

80

100

Percentages

20

40

60

80

100

Per centages

Fig. 1. Present and predicted mean proportions of densities with both projections: Markov and Stand level. a. Present and predicted
means considering only sexual regeneration in Moquinal. b. Present and predicted means considering sexual and a-sexual
regeneration in Moquinal. c. Present and predicted means considering only sexual regeneration in Aguirre. d. Present and predicted
means considering sexual and a-sexual regeneration in Aguirre. The mean is represented together with the standard deviation (in error
bars). Significant differences (p < 0.05) of the predictions with respect to the present values are indicated with an asterisk. Ea = Erica
arborea; Es = Erica scoparia; I = Ilex canariensis; Mf = Myrica faya; La = Laurus azorica; P = Prunus lusitanica; OS =Other species.

an increase or continuation of their importance in the

forest (Fig. 2c, e). Axis I discriminates between present
and projected plots. Axis II discriminates between shadetolerant and shade-intolerant species. Axis III explained
less than 3 % of the total variance and did not reveal any
relevant pattern in the distribution of the samples and
species.
The four projections were analysed for the present
plots with CCA (Fig. 3). The only explanatory variable
used in these analyses was present plots vs. projected
plots. As we have only one explanatory variable, the
analysis offered only one axis. Interpreted as a successional axis. Species with lower scores are projected to
decrease in importance, and those with higher scores are

projected to increase. Thus, these axes are useful for

comparing the behavior of different projections. In general, shade-intolerant species had low scores and L.
azorica had high scores, implying that L. azorica will
eventually replace shade-intolerant species. Prunus
lusitanica, depending on whether the projection considered the a-sexual regeneration or not, increased or decreased through the succession. We tested the eigenvalue
of each axis with the Monte Carlo test and all the
eigenvalues were significant (p < 0.05), though the significance needs to be interpreted with caution since the
observations are not independent.
When the two sites are analysed in a CCA (using a
dummy variable Moquinal vs. Aguirre) we find that

866

Arvalo, J.R. et al.

Fig. 2. First and second axis of DCA (eigenvalues were 0.307 and 0.204 respectively and the cumulative percentage variance of
species data of both axis was 61.1 %) for the eight present plots and the plots of the predictions (eight more per prediction). We
represented scores separately. a. Species scores. b. Present plot scores (p and the number of the plot) and predicted plots with Markov
projection from sexual regeneration. c. Present plot scores and predicted plots with Markov projection from sexual and a-sexual regeneration. d. Present plot scores and predicted plots with Stand level projection from sexual regeneration. e. Present plot scores and
predicted plots with Stand level projection from sexual and a-sexual regeneration. In graphs b-e the axes do not have the same scale.

the axis is significant (p < 0.05) for present species composition (eigenvalues and percentage of explained variance 0.203 and 50.6 % respectively), and significant for
stand projection including a-sexual reproduction (0.217
and 62.8 %). However, the axes were not significant in
any of the other three projections (percentage of explained variance of the axis less than 20 %). These
results, coupled with those of the DCAs, implies that asexual regeneration slows the successional convergence
between sites.

Discussion
Regeneration at the two sites was similar with Laurus
azorica dominating. The intermediate successional state
of this forest and the lack of frequent and large spatial
scale disturbance during the past 50 yr explain the similarities in sexual regeneration at both sites. The projections from both models using only sexual regeneration
indicated L. azorica and to a lesser degree other species
category as the most important species in the projected

plots (Fig. 2b, d). When a-sexual regeneration is considered, the predicted changes in species composition are
minor, indicated by shorter arrows of the projections
(Fig. 2c, e). Also Prunus lusitanica is predicted to increase its importance in the plots.
When only sexual regeneration is considered, both
models give similar results and predict a clear convergence between sites (Fig 2b, d). Some differences
emerged between both projections once the a-sexual
regeneration is considered in the models. The predicted
change in species composition is smaller for stand projection than for stand Markov projection (related with
the length of the arrow). Markov projection is a multigenerational model that predicts the species composition in a steady state while stand projection is a singlegenerational model. However, differences between the
projections were apparent only for two of the plots
(plots 2 and 8), while the general pattern of decrease of
shade-intolerant species and increase of shade-tolerant
species was maintained.
The stand projection method is computationally easier
and more straightforward. We considered it a useful

- Tree regeneration and future dynamics of the laurel forest on Tenerife Regeneration

867
Predictive model

a) Sexual-a-sexual

Stand level projection

b) Sexual

Stand level projection

c) Sexual-a-sexual

Markov projection

d) Sexual

Markov projection

Fig. 3. Species scores of CCA-axis I. Each horizontal line is the axis of a different analysis, each using present plot composition vs.
predicted plot composition as the explanatory variable. In the right of the graphs we specified the predictive model used. The
eigenvalues and the percentages of explained variance for these axes were: a. 0.110 and 19.7%. b. 0.236 and 35.0 %. c. 0.139 and
18.1%. d. 0.336 and 40.9 %. The Monte Carlo test indicated that each of the four axes were significant at p < 0.05. Ea = Erica
arborea; Es = Erica scoparia; I = Ilex canariensis; Mf = Myrica faya; La = Laurus azorica; P = Prunus lusitanica; OS =Other species.

model because summing saplings across the forest (stand

projection) gave similar results as allocating those saplings under particular canopy trees.
The effect of the incorporation of a-sexual regeneration
in the models is important for Prunus lusitanica and Ilex
canariensis. For these species their maintenance in the
forest canopy depend completely on the a-sexual regeneration. The results support the result of Condit et al. (1992):
A-sexual regeneration that tends to create aggregation is
responsible for the maintenance of some species. Also, asexual regeneration affects the dynamics of the forest by
slowing down the successional changes. Another effect of
the incorporation of a-sexual regeneration in the models is
the elimination of the clear convergence between sites.
We predict changes in the future canopy composition at both stations. Assuming little natural disturbance, we suggest that shade-intolerant species occupied some small patches inside the forest due to anthropogenic disturbances more than 50 yr ago, when the use
of the forest was more intense. Shade-intolerant species
are located in young patches that differ from the matrix
forest (at least a few generations old). Shade-intolerant
species are abundant in the seed bank (Arvalo 1998)
and disturbance may allow them to invade new space.
The seed bank is continually renewed by individuals at
the forest edge or corridors (very common in this forest)
where shade-intolerant species are dominant (Erica
arborea and Myrica faya) or from environmental patches
such as mountain peaks lashed by winds (Erica scoparia)
(Fernndez-Palacios & Arvalo 1998). Unpredictable large
disturbances may cause a reversion in the community
composition. If disturbance is reduced, changes will be
determined by biological relationships and habitat

availability (Franklin et al. 1993).

The results of multivariate analysis are easier to
understand and interpret than the comparison of species
individually. With a low sample size it has been possible
to detect an interpretable difference between present
and projected plots, site of the plots and species groups
(shade-tolerant and -intolerant species). However,
multivariate analysis could be especially useful in cases
of large data sets, where trends of species abundances
are not easy to extract from observation. In addition,
ordination gives us insight into the directionality of
change in the forest.
We chose two representative sites of the laurel forest
of Tenerife and selected the plots randomly. The results
could be extrapolated to similar laurel forests of Tenerife,
although a high variability in composition and structure
has been recognized in these forests (Santos 1990).
In order to understand the dynamics of the laurel
forest more completely, we recognize the necessity of a
long-term study with representative permanent plots
(van der Maarel 1993) where it is possible to empirically
test the projections and to examine the dynamics of the
laurel forest in finer detail.
Acknowledgements. This work is part of a research project in
laurel-forest dynamics (No. 95 6129 05) supported by the
Canarian Government (Consejera de Poltica Territorial) and
European Union (LIFE projects program). We thank Sue
McAlister, Steven Thompson and the members of the ecology
group of the Department of Botany (Oklahoma State University) for advice on the data analysis and help with the manuscript. We also thank Peter S. White and three reviewers for
their constructive comments.

868

Arvalo, J.R. et al.

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Received 24 November 1997;

Revision received 20 June 1998;
Final revision received 8 March 1999;
Accepted 8 March 1999.