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1/14
Journal f Tropical
Ecology (1990)
6:307-320. With
figures
Preliminary
studies on
forest
tructure nd
floristicson Volcan Barva, Costa Rica
ANGELA
HEANEY and
JOHN PROCTOR
Department
of Biological
and Molecular
Sciences,
University
f Stirling, tirling
FK9
4LA,
Scotland
ABSTRACT. Volcin
Barva,
Costa
Rica,
has on
its
northern
lope
an
unbroken
equence
of
rain
forest n
volcanic
parent materials
romnear sea
level
at La
Selva Field
Stationup
to its sum-
mit at
2906
m. It provides good
area to study
forest hangeswith
ltitude nd their auses.
In
the
present paper we
describe the forests s a
backgroundfor soil and
litterfall tudiesfrom
1
ha plots
at each of the
following ltitudes: 100 m, 500
m, 1000 m, 1500 m,
2000 m and
2600 m. The
canopy
heights with heightof the
highest
mergent
n
parentheses)
anged
rom
35-40 m
(45 m) at 100 m to
20-23 m (32
m) at 2600 m; basal area
was least (22.7 m2)
at
1
00
m
and highest 51.2
m2)
at
2600 m; the tree
>10
cm
dbh) density angedfrom
91 ha-'
at 500
mto 617 ha-' at
2600 m. Most treeswere
dentified nd on samples
of themwe
recorded
presenceof buttresses,
ianes, kiophytic
limbers, ascular piphytes nd
bryophytes; nd drew
profilediagrams. n the classification f Whitmore 1984) the two lower plots are evergreen
lowland
rain forests; he other four
are lower montanerain
forest.
pecies richnesswas highest
in
the
plot at 500 m,with at least
135 species of tree, nd
least at 2600 m, with at
least 35
species. The
Volcin
Barva forestaltitudinal
sequence is briefly
ompared with those else-
where.
KEY
WORDS: altitudinal
onation,Costa Rica,
lowland rainforest,
montanerainforest.
INTRODUCTION
There
have
been several studies of
forest
onation
on
single
tropical
inountains,
e.g.
Brown
(1919)
for Mount
Maquiling
in
the
Philippines,
Grubb &
Stevens
(1985) in Papua New Guinea, Proctor et al. (1988) in Sabah, and Whitmore
(1972)
and
Whitmore
& Burnham
1969)
in
Malaya. Apart
from
Beard's
(1944,
1946,
1949) descriptions of
small mountains in
the
Caribbean such
studies
were
apparently
lacking
for the Central
American
tropics.
In
1985
the
oppor-
tunity
arose
to
work on Volcain
Barva,
Costa Rica, which
has
on its northern
slope,
an
unbroken
sequence of rain forest
from
near
sea
level
up
to
its summit
at
2906
m
(Figure
1). The
forests n the lower part of this
sequence
include the
the
area
around La
Selva Field
Station
whilst
the
upper
part
is within
the
Braulio
Carillo
National Park. The Barva
vegetation
has
been
briefly
described
in a general way by Hartshorn & Peralta (1987). Our aim in thepresentpaper
is
to
provide
preliminary
descriptions
of
six plots
as
a
background
for studies
(307)
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308 ANGELA HEANEY AND
JOHN
PROCTOR
Puerto
Viejo.
*
a
Selva
Field
Station
500
m
1500m
P
Vcan
Cacho
Negro
2000
m
A
N
2600
m
A&
VoIcan
Barva
2900
m
o
5
10 km
l
l
l
N
ICARAGUA
COSTA
RIC
Figure
1. The locations
ofthe
study
plots
on
Volcin
Barva,
Costa Rica.
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Forest on
Volcan
Barva
309
on
their
soils
(Atkin & Proctor
1988, Grieve
et al. 1990, Marrs
et
al.
1988)
and litterfall
Heaney &
Proctor 1989).
THE STUDY PLOTS
The study was carried out
along
a
transect
fromthe
La
Selva
Biological
Station
(100 24' N,
840
00'
W) up
the
northern
lope
of Volcain Barva.
One
plot
of 1
ha
of
largely
mature-phase
forest
was investigated
at each
of six altitudes:
100,
500, 1000,
1500, 2000
and 2600 m. The plots were
permanently
marked
and
each
sub-divided nto
twenty-five 0
X
20
m
sub-plots. The
locations are
shown
in
Figure
1
and a
summary
of
some features
of
the
topography
including
the
slope) and
vegetation of the six plots is
given n Table 1.
At La Selva the mean annual rainfall (1957-1979) was 4210 mm and the
mean annual
temperature
240C. The
rainfall s
relatively
aseasonal
(Figure
2).
Rainfall
data
are
not
available
along
the
transect,
but
data
collected
from
elec-
ted
stationsnearby (Table
2) suggest that it may be
greatest
n the
mid-region
of
the mountain. The
temperature apse rate
is
not known.
Ground
frostswere
observed
in
a
clearing at
2600 m on
several days
in
April
1985,
but not
during
continuous
daily observations between 5
March and 5 April 1985
in a
clearing
at 1800 m.
Table 1. Some
details of the
1
ha study-plots nd trees
> 10 cm dbh) at a range f altitudes
n Volcin
Barva,Costa Rica.
Tallest
No. of
No.
of
Plot
Basal
Canopy
tree No. of missing
uniden-
altitude
Slope area
height
height
indi-
No. of speci-
tified
(m)
(0)
Aspect
(Mi2) (m) (m) viduals
species mens trees
100
7
E
22.7 35-40
45
494
111
1
4
500
7
N
24.3 30-35
50
391
135
6
4
1000 7
NE
31.2 30-35
45
546 109 12 6
1500 7
NE
29.2 25-30
38
553
65 15 39
2000
10
NW
28.6
20-25
35
448
69
13
19
2600
15 N
51.2
20-23
32 617
35
1 35
Table
2. Mean annual rainfall or fiverainfall tations
t a
range
of
altitudes
near the
forest
tudyplots
(from
Hartshorn Peralta
1987).
Distance
km)
Altitude
f
and direction
nearest Duration
of
Altitude
from
nearest
studyplot
Rainfall observations
Name
(m)
study plot
(m) (m)
(yrs)
La
Selva
42 3, N
100 4015
26
San
Miguel
500
11,
W
500 4627
17
Cariblanco 970 10,W 1000 5096 5
Vara
Blanca
1804
5, W 2000
3426 21
Sacramento
2260
8, S 2600
3268 11
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310
ANGELA HEANEY AND JOHN PROCTOR
1200
1000
800L
0
E
E
E600-
400-
200
-
0
J F
M A
M
J
J A S
0
N D
Time (calendar month)
Figure 2.
Mean annual
rainfall or
1
October 1957
-
31 December 1979
for
La
Selva Field
Station,
Costa
Rica.
The vertical
bars
indicate
maxima and
minima
for
each
month
during
the
measurement
eriod
(La Selva Field Station unpublished).
Some aspects of the soils have been described by Marrset al. (1988). The
parent materials are basaltic and andesitic lavas of Plio-Pleistocene age with
a
trend towards tuff or agglomerate-likematerials with increasing levation.
The
soils all have low concentrations of available phosphorus and exchangeable
bases with no
clear altitudinal trends. The highest
concentrations were
in
the
plot
at
2600 m. Rates of nitrogen mineralization and nitrification
howed
a
clear
trend of decrease with altitude.
MATERIALS AND
METHODS
The
six
1
ha
plots were measured out without slope correction. The 100 m
altitude was judged from a map, the others were estimated using
an altimeter.
Five
of
the plots were 100
X
100 m but that at 2000 m was of an irregular
shape
to
avoid a ravine which was about 3 m away from the south-western
edge
of
the
plot.
All
trees
(>10
cm
dbh) were enumerated and
their
diameters measured,
usually
at
breast
height (1.3 m) except for those with buttressesor prop
roots
which
were
measured 30 cm
above
the protrusion's unction
with
the bole.
Some
trees had multiple stems (>10 cm dbh) and each stem was
measured
separately. A transect (60 X 7.5 m), which had a substantial proportion of
mature
forestwas
selected
for
a profile diagram of trees >6 m high).
At
each
site,
a
sample of 100 trees 80
at
2600 m)
was
selected
in
their
order
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Forest on
Volcan
Barva 311
of enumeration from a random
point.
For these selected trees the
following
were
recorded: the presence or absence of contacts with photophytic climbers
(lianes) in three size categories, skiophytic climbers (defined by Grubb
et
al.,
1963), and vascular epiphytes; and the percentage cover of epiphytic mosses
on the bole above the ground at 2 m. The trees have mostly been identified
(by
G.
S. Hartshorn) at least to family evel and separate taxa recognized
to
allow construction of species-area curves. The number of missing specimens
and wholly unidentified trees (which were not included in the species-area
curves) in each plot is given n Table 1.
RESULTS
The profile diagrams (Figures 3-8) give an impression of the overall appearance
of the
forests nd show the changes in stature with altitude.
Some
forest structural characteristics are given in Tables 3-5. The
most
consistent altitudinal
trend is the decrease
in
tree height Table 1).
The
plot
at
2600
m has a
high
basal area
(51.2
m2)
compared
with
the lower
plots (Table
1). The relatively small differencesbetween the plots for percentage of trees
45 _
40,
Ho
Figure 3. Profilediagram 60 X 7.5 m) of forest t
10)0
m on the
Volcain
Barva transect, osta Rica.
Trees
less than 6 m high excluded. Symbolsfor identified rees over 10 cm dbh: Bi, Brosimum
actescens
(Moore) Berg; Ca, Casearia arborea Rich.) Urban; Em, Euterpe
macrospadixOerst;Ga, Guatteria erugi-
nosa Standl.; Gr,
G'uarea
rhopalocarpa Radlk.; Gx, Guarea sp.; Ho, Hieronymaoblonga var.
benthamii
(Tul.)
Muell.
Arg.; 1,
nga longispicaStandl.; Ip, I. punctata
Willd.;
Lix, Licania sp.; Mm,Miconia
multi-
spicata Naud.; Nn, Naucleopsis naga Pittier; a, Pourouma aspera Trecul; Pc, Protium ostaricenseRose)
Engler; Pg, P. glabrum Rose) Engler;
Pm, Pentaclethramacroloba
(Willd.)
Kuntze; Pp, Protium
pana-
mense (Rose) I. M. Johnston;Qb, Quararibea bracteolosa (Ducke) Cuatr.; Vh, Vochysia
hondurensis
Spragne;Wg,Welfia eorgiiWendl.
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6/14
312 ANGELA HEANEY
AND
JOHN
PROCTOR
40
O-
Figure 4. Profilediagram 60 X 7.5 m) of forest t 500 m on the
Volcain
Barva transect, osta Rica. Trees
less than 6 m high excluded. Symbols for dentified rees over 10 cm dbh: Ax, Ardisia sp.; Cp, Couepia
polyandra (Kunth.) Rose; Cs, Colubrina spinosa Donn. Smith;
Dm,
Dussia macroprophyllataD.Sm.)
Harms; Dp, Dystovomita
ittieri
Engl. W.G. D'Arcy; Gy, Guarea sp.; Ig, riartea igantea
Wendl.;
t, nga
thibaudiana D.C.; Lax, Lauraceae sp.; Mc, Macrolobium costaricenseW. Burger;Pac, Parathesis hryso-
phylla Lundell; Rx, Rubiaceae sp.; Ry, Rubiaceae sp.; Vf, Vochysia errugineaMart.
40
E~~ ' v
0A0.
35
-
~
W~-
30(-
k
V
25
-j~
20
15~
5
0-
Figure 5. Profile diagram 60 X 7.5 m) of forest t 1000
m
on
the
Volcan Barva transect,
osta
Rica.
Trees
less
than 6
m
high excluded. Symbols for dentified rees over
10
cm dbh:
Bic,
Billia
colombiana
Planch
&
Lind.; Ca,
Casearia arborea (Rich.) Urban; Ce, Cassipourea lliptica Poir.; Dp, Dystovomita
pittieriEngl.W.G. D'Arcy; Em, Euterpe macrospadixOerst.;Gg,Guareagrandifolia C; Hx,Humiriaceae
sp.; Ix, Inga
sp.;
Lay,
Lauraceae sp.; Liy, Licania sp.; Ny, Neea sp.; Nx, Nephelea sp.; Po,Pseudolmedia
oxyphyllariaDonn.
Smith; Rg, Rubiaceae sp.; Rh, Rubiaceae sp.; Sx, Sapotaceae sp.; Vf, Vochysia
ferruginea.
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Forest on
Volcan
Barva
313
35
30
Pam
~
~~~~~~~~~~)~~~~~~~~~~~~~~~~~~~~~~~~~-
15-
0
Figure
6.
Profile diagram 60 X
7.5 m) of forest t 1500
m on the VolcainBarva transect, osta Rica.
Trees less
than6 m high xcluded.
Symbolsfor dentified
reesover 10 cm dbh: Bh, Billia
hippocastanum
Peyritsch; x, Clethra p.; Cz,
Cyatheaceae; Da, Dendropanax arboreus
L.) Dcne. & Planch.; Dw, Drimys
winteri
orst.; Ea,
Elaegia
auriculataHemsl.; Ex,
Eugenia storkii tandl.;
Go, Guatteria liviformis onn.
Sm.; Hp, Hieronymapoasana
Standl.; II, Inga longispica Standl.;
Laz, Lauraceae sp.; Mg, Matayba
sp.;
Mam,
Macrohasseltia
macroterantha
Standl.
&
L.
Wms.)
L.
Wms.;
Ox,
Ossaea
sp.; Pad,
Parathesis
denan-
theraHook.
f.; Pam,Persea americanaMill.; Rf,
Rapanea ferrugineaR. & P.)
Mez.
30
Figure 7. Profilediagram 60 X 7.5
mn)
of forest t 2000 m on the
Volcain
Barva transect,Costa Rica.
Trees less than 6 m high excluded. Symbols for
dentified rees over 10 cm dbh: Ap, Ardisiapalmana
Donn. Smith; Bc, Brunelliacostaricensis tandl.; Cw, Cinclima p.; Cy, Cyatheaceae;
Dip,
Didymopanax
pittieriMarch.; Hy, Hamamelidaceaesp.; Hp, Hieronyma oasana
Standl.; Mx, Miconia sp.; My,Miconia
sp.; Mz, Melastomataceae p.; Px, Piper sp.; To,
Tu4rpinia
ccidentalis SW.) Don; Tx,
Tetrorchidiu4m
p.;
Vm,
Vibu4rnum
exicanum nomen).
in different iameter classes (Table
3) is remarkable. The frequency of buttres-
sed trees remains relativelyhigh with increasingplot altitude (Table
4)
although
the actual numbersof tall (>1 m high) buttresseson the sampled treesdecreases
markedly: there were 22 at 100 m,
39 at 500 m, nine at 1000 m, three at
1500 m, four at 2000 m, and five at
2600 m. Lianes decreased in
numbers
with
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314 ANGELA HEANEY AND JOHN PROCTOR
30-
20-
CtA\t2
I~~~~~~~~~~~~~~~~~~~~~I
oL
Figure 8.
Profile
diagram 60
X 7.5
m)
of
forest
t
2600
m
on the
Volcin Barva
transect,
osta Rica.
Trees less
than
6 m
high excluded. Symbols for identified
rees
over
10 cm dbh:
Ay, Ardisia sp.; Bc,
Brunellia costaricensis tandl.; Cx, Clethra sp.; Dx, Dendropanax sp.; Iv, Ilex vulcanicola Stand.; Vm,
Viburnum
mexicanum nomen).
Table
3.
Percentages
f trees
>
10
cm dbh) in
a
range
of
diameter-classesn six plots on Volcan Barva,
Costa Rica.
Plot
Diameter-class
cm)
altitude
(m)
10-20
20-30 30-40 40-50 50-60
60-70
70-80 80-90
90-100
>100
100
66.7
17.0 6.2 3.1 3.1
1.8
0.6
0.2
-
0.2
500 57.8 19.8 11.6
3.6 3.1
1.0
1.5
0.2
0.2 1.0
1000
59.6
18.5 8.9
6.0
2.7
1.3 1.3
1.3 0.2 0.2
1500
51.4 24.5
13.4
6.5
1.8
0.9
-
- -
1.4
2000
44.1
26.6 16.9
5.2
2.5
2.0 0.7
-
0.2
1.8
2600
51.4
22.8
8.5
7.2
4.6
2.0 1.0
0.6
0.3
1.6
Table 4. The percentageof sampled trees N
=
100 except for the plot at 2600 m where N
=
80) with
buttresses, ianes, skiophytic limbers, ascular piphytes, nd with more than 50% of
the boles covered
by bryophytes
t
2
m
from
he
ground, n plots at a range f altitudes n Volcin Barva,
Costa Rica.
Buttresses
Lianes
Plot of heights cm) of diameter cm) Bryophytes
altitude
Skiophytic Vascular (>50% cover
(m)
50-100
>100
>1-5 >5-10
>10 climbers
epiphytes
at
2
m)
100
23
6
51
40
2
75
89
5
500
23
8 45
25
7
82 86
4
1000
17
5
33 12 2
71
92 32
1500 7
3
18 7 0 95
99
79
2000 14 3 7 4 0 85 100 72
2600
16
5
14
0 0
18
100
80
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Forest on
Volca'n
Barva
315
140-
130- 0
O Om
50
m
120-
0
l
OOOm
A 1500m
110- *
2
OOOmX
A
2600m
100
5
o
90-
*
80-
.0
70-
E
560-
u
50-
cn
40-
30-
20-
10-
5 1
1'5
20
25
Sub-plots (number
of 20x20m)
Figure9. Species-area curves
for trees > 10 cm dbh) on the six plots on Volcin
Barva. The numbers f
missing pecimens
and
unidentified rees not included in these curves
s
given
n Table
1).
Each
curve
follows
he
orderof enumeration
f the twenty-five0 X 20 m sub-plots.
altitude and
none >5
cm
diameter were recorded
from
the
highestplot. Vascu-
lar
epiphytes were abundant at all
altitudes whilst bryophyte cover increased
markedly
with
altitude.
There
is
a
trend
of
reduction of species richness with altitude
(Table
1
and
Figure 9) although the
plot
at 500
m is much richer than
that at 100
m.
An
increase
in
the number of
missing or unidentified pecimens
partially accounts
for
the drop
in
species numbers between 1000
and
1500 m.
Above 500
m
there
is
a trend of decreasing
species numbers and the plot
at
2600
m
is
by
far
the
most
species poor.
Table 5 shows dramatic changes in tree family composition with altitude.
At 100 m
there is a
high proportion
(32.9%
of
the
basal
area)
of the
Mimo-
saceae
of
which most
(28%
of
the basal
area)
is
contributed
by
Pentaclethra
macroloba. The
Mimosaceae
are
still
the
leading family (11.9%
of the basal
area)
at
500 m
although Pentaclethra
macroloba
is
absent.
At
1000 m the
Meliaceae
(10.4%)
and the
Mimosaceae
(10.1%)
are commonest
and
similar
n
their
proportion of the
basal area, whilst the Euphorbiaceae have the
highest
percentage
of
basal area at
1500 m (14.5%)
and
2000
m
(21.7%).
Tree
ferns
(Cyatheaceae)
make a
substantial contribution
to the
basal area
at 1500 m
(9.0%)
and
2000
m
(10.6%). The plot at 2600 m has a
preponderance
of
Araliaceae (25.7%) and Aquifoliaceae (21.9%) whilst oddly, in view of their
abundance
at 2000 m, the Euphorbiaceae are
absent.
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316
ANGELA HEANEY
AND
JOHN
PROCTOR
Table 5. The tree familieswith their
percentage
f total basal area and theirnumbers
f ndividual
rees
(>
10
cm dbh)
in
the six studyplots at a range f altitudes n Volcin
Barva,Costa Rica.
Plot altitudes
100
m 500 m 1000
m 1500 m 2000 m
2600 m
BA
(%) No.
BA
(%) No. BA (%) No. BA
(%)
No.
BA
(%)
No.
BA
(%)
No.
Actinidiaceae 0.48
2
0.03
1
Anacardiaceae 2.15
4
3.09
4
Annonaceae
1.42
30
3.82
15 0.58
5
4.80
29
Apocynaceae 0.23
4
Aquifoliaceae 0.06
1 21.91 72
Araliaceae
3.33
14
3.09
10
0.81
3
5.40
25
3.91
16 25.68
134
Betulaceae
2.32
4
Bignoniaceae
0.14 1
Bombacaceae 0.28 4 0.24 2
Boraginaceae 0.84 5 0.03
1
Brunelliaceae
0.35 2 3.97 15 6.87
65
Burseraceae 7.29 47 4.30 28
Caesalpiniaceae 0.99 9
4.06 15
Capparidaceae 0.05
2
Caprifoliaceae
4.69 25 0.63
7
10.74
128
Caricaceae
0.05
1
Celastraceae 1.10
5
0.12
1
Chloranthaceae
0.43
7 0.03
1
Chrysobalanaceae
0.04
1
5.77 17 3.63
13
Clethraceae
0.13
2
0.26
2
3.75 18
Clusiaceae
0.38 2 0.49
1
0.16
4
Combretaceae
0.56 2 0.64
2
Compositae 0.31 1 0.62 19
Coinaceae
1.37 3
Cunoniaceae
0.29
2 1.23
5
12.75
12
Cyatheaceae
3.01 50 8.96
73
10.59
60
1.74
35
Dichapetalaceae
0.04 1
1.22
3
Dilleniaceae 0.45 9
0.04
1
Elaeocarpaceae 0.08 1
0.54
6
0.39
1
Erythroxylaceae 0.04 1
0.85
7
Euphorbiaceae 4.0
10
5.40
10
3.98
13
14.50
63
21.74
60
Fabaceae 1.38 12 5.82 9
5.18 17
Fagaceae 1.68
4
Flacourtiaceae
2.16
11
1.82
11
2.92
15
2.67
9
Guttiferae
0.16
2
0.78
9
6.39 26 8.49
35
0.48
3
Hamamelidaceae
6.48 24
Hernandiaceae 1.81 5
Hippocastanaceae
2.61 3 5.64 21 3.25
14
2.34
4
0.06
1
Humiriaceae 1.25
1
0.05
1
2.65
7
Icacinaceae
0.57
10
1.21
15
Lacistemataceae 0.05 1
0.65
6 0.06
1
Lauraceae 1.14 11
1.64
14
3.90
14
5.82
19
4.76
22
4.33
32
Lecythidaceae
0.58 1
1.18
2
Malpighiaceae
1.23 2
0.69
2
Malvaceae
0.06
1
0.12
1 0.06
1
Margraviaceae
0.04
2
Melastomataceae 0.10
2 1.64
10
1.12
13
1.10
16 6.83
61
0.28
11
Meliaceae 3.90
17
6.92 32 10.38 52 0.29
7
0.78
6
Mimosaceae
32.94
69 11.85
40
10.09
33
6.99
35
Moraceae
5.82
35 3.34
11
2.66
21
0.08
2
Myristicaceae 1.76 8 2.23 5 0.53 6
Myrsinaceae 0.14 3 2.20 13
0.33
3
1.70 24 5.10
39
3.24
36
Myrtaceae
0.14
2 0.28 2
1.03 24
1.00
18
1.30
7
Nyctaginaceae
1.67 21 0.36
1
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Forest on Volcan Barva
317
Table 5
-
continued
Plotaltitudes
100 m 500 m 1000 m 1500 m 2000 m 2600 m
BA
(%) No.
BA
(%)
No.
BA
(%)
No.
BA
(%)
No. BA
(%)
No.
BA
(%)
No.
Olacaceae 0.99
5 2.27 3
Palmae 10.02 114 0.57 7 3.40 55
0.07 2
Piperaceae
0.04 1 0.52 11 0.02 1
Polygonaceae
0.33 1
Quiinaceae 0.07 1 0.06 1
Rhamnaceae 0.14
2 0.40 3
Rhizophoraceae 0.12 2 0.11 2 0.16 2
Rosaceae
0.76 7 0.17 1 0.08 1
Rubiaceae 0.99 11 2.51 50 2.88 47 3.00 30
2.70 8
Sabiaceae
0.36 2 2.07 6
2.87 13
Sapindaceae 0.09 4 0.71 3 0.05 1 0.35 4
0.27 2
Sapotaceae 2.82 9 3.15 8 7.08 17 3.35 10
Simaroubaceae 0.94
2
Solanaceae
0.70 8 0.11
4
Staphyleaceae
2.29 3 6.14 18
Sterculiaceae
0.18
2 0.30 2
Styracaceae 0.04 1
Symplocaceae
0.14 1
0.49 3 2.41 10 0.18 1
Theaceae
3.27 1 0.09 11 0.58 3 0.11 1
Tiliaceae
3.66 10 1.82 3 0.21 2
Verbenaceae
3.35
1
0.52 3
Vochysiaceae 5.10
2 4.66 5 11.27 10
Winteraceae 0.23 2
2.28 7 0.11 1
Unidentified 0.05 5 0.89 10 3.16 18 14.97 54 7.76 32 4.9 36
or missing
specimens
DISCUSSION
Forest
types
Further nformation
on the
La
Selva
forests and
those
of
the
Barva
transect
is
in
Frankie
et
al.
(1974),
Hartshorn
(1983),
Hartshom & Peralta
(1987)
and Heaney (1988). In the classificationsystemof Whitmore 1984) the two
lower plots
are
probably evergreen owland
rain forest
and the
other four
are
lower montane rain forest although we lack the
leaf-size information
which
is
the most
objective
criterion
for
the classification.
From
Hartshorn
&
Peralta
(1987)
the
plots
are said
to
fall
nto
the
following
Life Zones
(Holdridge
et
al.
1971):
100
m, Tropical
wet;
500
m, Tropical wet,
cool
transition;
1000
m, Premontane rain; 1500
and
2000 m,
Lower
montane
rain; 2600 m,
Montane
rain.
The Life
Zone classification
system
remains
vague,
however,
and
objective
definitions
n terms
of
structure
nd
physiognomy
are
lacking.
The lowland forestsat 100 m and 500 m have a tall statureand are species
rich
by
Central
American
standards
(Hartshorn 1983, Holdridge
et
al.
1971).
The forest of
Corcovado,
Costa
Rica
which
has
100-120
tree
species
haI
was
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12/14
318 ANGELA HEANEY
AND JOHN PROCTOR
described by Hartshorn (1983) as
'undoubtedly
the
richest forest in Central
America'. The plots described in the
present paper
at
100, 500
and 1000
m
which have at least 111, 135 and 109 species respectively,
require
this state-
mentto be reassessed.
It is beyond the scope of this paper to review altitudinal changes
in
wet
tropical forests n general. Many
of
the
trends in structure
and
floristics on
Volcain
Barva
(Tables 1, 3,
4 and 5 and
Figures 3-10)
could be
predicted from
other studies (e.g. Grubb 1977) although
regrettablywe
have no information
on
leaf size and shape, frequency of
pinnate leaves, and frequency
of cauli-
flory. The most surprising features on
Volcan
Barva were perhaps that the
largest tature and most species-rich lot was at 500 m ratherthan at the lowest
altitude 100 m) and that the greatest
basal area occurred in the highestplot.
In
view of the recent paper on litterfallHeaney & Proctor 1989) it is
useful
here to compare the stature of the Volcain Barva forests with that of other
montane forestswhere production and
nutrient yclinghave been studied.
The
plots at 1500 m and above on
Volcain
Barva have taller stature trees than:
Jamaican
forests
at
1550
m
(with canopy trees up
to
18
m
tall) (Grubb
&
Tanner
1976); Gunung Mulu, Sarawak
(where
the tallest
trees were 21
m
high
in a
plot at 1310 m, 15 m at 1860 and 5
m near the summit at 2340 m) (Martin
1977); and Gunung Silam, Sabah
(where above 700 m the tallest trees were
21
m) (Proctor
et al.
1988).
Taller
montane forests n which the trees
exceed
the
height of those in the two upper
Volcain
Barva plots are those at ca. 2500
m
on Mount Kerigomna, Papua New Guinea, which have a canopy of 27-33 m
with
emergents p to 37
m
(Edwards
&
Grubb 1977); and at Merida, Venezuela
at
about 2300 m with a canopy
height of 35-40 m (Grimm & Fassbender
1981).
At
present
there is
no unifyinghypothesiswhich satisfactorily xplains
the
altitudinal changes of the forests on
Volcain
Barva and the differences etween
these
forests nd those elsewhere.The likelihood that decreasing nitrogen upply
with altitude
is
important
on Volcain Barva
is
discussed
by Heaney
& Proctor
(1989)
and
Marrs
et
al. (1988). Generalizations fromecological work
on Volcan
Barva
are
hindered because of the
non-uniformity f the
mountain's
lithology
and its lack of really low stature upper montane or sub-alpine forest.The
answers
to
many of the questions about forests of wet tropical mountains will
be facilitated
by long-term nd experimental studies. In this respect the Volcain
Barva forests
are
ideal since they now have a fully protected status and are
close
io
the excellent facilities of the
La Selva Field Station. The six plots
discussed
here and others set up later are
currently eing monitored for growth
and
regeneration
studies
by
D. and
M.
Lieberman
who
are also
obtaining
more
detailed
climatic
nformationfromthe mountain.
ACKNOWLEDG
EMENTS
We thank
National
Parks
Service of Costa Rica forpermission to work in Braulio
Carillo,
The
Organization for Tropical
Studies for permission to work in the
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13/14
Forest
on
Volcan
Barva
319
grounds of
La
Selva Field Station, Operation Raleigh for
support,
its staff
(particularly Mr K.
Hamylton-Jones) and venturers for help, and Dr G. S.
Hartshorn for help
in
the
field and
for his tree
dentifications.Drs P.
J.
Grubb,
D. L. Kelly and E. V. J. Tanner are thanked forcomments on themanuscript.
The
work was supported financially by the British Ecological
Society, the
Carnegie Tru'st, the Leverhulme Trust and the National
Environmental Re-
search Council.
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DDT and its deriviates
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environmental aspects. Environmental Health Criteria, No.
83.
World Health Organization. 1989. 98 pages. ISBN 92-4-154283-7. Price:
Sw. fr.
13.-;
US$ 10.40. Order no. 1160083.
This small book considers the effects of DDT and its metabolites on populations of organisms
in the environment from an ecotoxological point of view. The opening sections deal with the
properties of these substances that help to explain their resistance to degradation, their
widespread persistance in the environment, and their high potential for bio-accumulation.
The major part of the book is concerned with their toxicity to micro-organisms, and aquatic
and terrestrial animals, with the emphasis on fish and birds. Ecological effects from field
application are also discussed. Numerous controlled laboratory experiments documenting
direct and sublethal effects are cited (20 pages of references). In the final evaluation, the
book
concludes that these compounds should be regarded as a major environmental hazard.
Obtainable from: World Health Organization, Distribution and Sales, 1211 Geneva 27,
Switzerland.
KARTIKASARI,
S. N.
&
WHITTEN, A. J. 1989. The natural resources, ecology and
environ-
ment of Java and Bali: a
bibliography.
(in English
and
Indonesian). EMDI, Jarkarta.
320
pages. ISBN 979-8115-00-7.
Obtainable from: Publications Office, EMDI, Kantor Menteri Negara, Kepen Kependudukan
dan Lingkungan Hidup, Jalan Merdeka Barat 15 Jakarta 10110. Indonesia.
ZEIN AHMED
ZEIN &
HELMUT KLOOS, (eds). 1988.
The
ecology of
health
and disease
in
Ethiopia. Ministry of Health, Addis Ababa.
xi
+
319 pages.
This
book reflects
the main
concerns
and
issues
in
health
development
in
Ethiopia today.
Adopting
an
ecological approach,
it identifies
and
suggests
solutions
for some
of
the
major
health problems
of
the population. Contributors are
from various fields
in
public
health
and
the
medical and social sciences, and it
is
intended
for
Ethiopian graduate
and
undergraduate
students
in
these disciplines, as well as medical and public health researchers, planners
and
policy-makers.
Obtainable
from: Ministry
of
Health, Planning,
and
Programming Bureau, through:
Zein
Ahmed, PO Box 109, Gonder, Ethiopia, and Helmut Kloos, PO Box 31609, Addis Ababa,
Ethiopia.
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