Journalof TropicalEcology (1990) 6:307-320.

With9 figures

Preliminary studies on foreststructureand

floristicson Volcan Barva, Costa Rica
ANGELA HEANEY and JOHN PROCTOR
Department of Biological and Molecular Sciences,
Universityof Stirling,StirlingFK9 4LA, Scotland

ABSTRACT. Volcin Barva, Costa Rica, has on its northernslope an unbrokensequence of rain
foreston volcanic parentmaterialsfromnear sea level at La Selva Field Stationup to its summit at 2906 m. It providesa good area to studyforestchangeswithaltitudeand theircauses. In
the presentpaper we describe the forestsas a backgroundfor soil and litterfallstudiesfrom
1 ha plots at each of the followingaltitudes: 100 m, 500 m, 1000 m, 1500 m, 2000 m and
2600 m. The canopy heights(with heightof the highestemergentin parentheses)rangedfrom
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
100 m and highest(51.2 m2) at 2600 m; the tree (>10 cm dbh) densityrangedfrom391 ha-'
at 500 m to 617 ha-' at 2600 m. Most treeswereidentifiedand on samplesof themwe recorded
presenceof buttresses,lianes,skiophyticclimbers,vascularepiphytesand bryophytes;and drew
profilediagrams.In the classificationof Whitmore(1984) the two lower plots are evergreen
lowland rain forests;the other fourare lower montanerainforest.Species richnesswas highest
in the plot at 500 m, with at least 135 species of tree,and least at 2600 m, with at least 35
species. The Volcin Barva forest altitudinal sequence is brieflycompared with those elsewhere.
KEY WORDS: altitudinalzonation,Costa Rica, lowland rainforest,montanerainforest.
INTRODUCTION

There have been several studies of forestzonation 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 fromBeard's (1944,
1946, 1949) descriptions of small mountains in the Caribbean such studies
were apparently lacking for the Central American tropics. In 1985 the opportunity arose to work on Volcain Barva, Costa Rica, which has on its northern
slope, an unbroken sequence of rain forestfromnear sea level up to its summit
at 2906 m (Figure 1). The forestsin 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 brieflydescribed
in a general way by Hartshorn & Peralta (1987). Our aim in the presentpaper
is to provide preliminarydescriptions of six plots as a background for studies
(307)

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308

ANGELA HEANEY

AND JOHN PROCTOR

PuertoViejo.
*

a Selva Field Station

500 m

1500m
Vcan
2000 m

Cacho Negro

2600 m
A&

VoIcan Barva
o
2900 m

10 km
l

N ICARAGUA

COSTA RIC

Figure1. The locations of the studyplots 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 transectfromthe La Selva Biological Station
(100 24' N, 840 00' W) up the northernslope 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 permanentlymarked and
each sub-dividedinto twenty-five20 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 givenin Table 1.
At La Selva the mean annual rainfall (1957-1979) was 4210 mm and the
mean annual temperature 240C. The rainfall is relativelyaseasonal (Figure 2).
Rainfall data are not available along the transect,but data collected fromselected stations nearby (Table 2) suggest that it may be greatestin the mid-region
of the mountain. The temperaturelapse 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-plotsand trees (> 10 cm dbh) at a rangeof altitudeson Volcin
Barva,Costa Rica.

Plot
altitude
(m)

Slope

100
500
1000
1500
2000
2600

(0)

Aspect

7
7
7
7
10
15

E
N
NE
NE
NW
N

Basal
area
(Mi2)

22.7
24.3
31.2
29.2
28.6
51.2

Canopy
height

Tallest
tree
height

(m)

(m)

35-40
30-35
30-35
25-30
20-25
20-23

45
50
45
38
35
32

No. of
indi-

No. of

viduals

species

494
391
546
553
448
617

111
135
109
65
69
35

No. of
No. of
missing unidenspecitified
mens

trees

1
6
12
15
13
1

4
4
6
39
19
35

Table 2. Mean annual rainfallfor fiverainfallstationsat a range of altitudesnear the foreststudyplots

(fromHartshorn& Peralta 1987).

Name

La Selva
San Miguel
Cariblanco
Vara Blanca
Sacramento

Altitude
(m)

42
500
970
1804
2260

Distance (km)
and direction
fromnearest
study plot

3, N
11, W
10, W
5, W
8, S

Altitudeof
nearest
studyplot
(m)

100
500
1000
2000
2600

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Rainfall
(m)

4015
4627
5096
3426
3268

Durationof
observations
(yrs)

26
17
5
21
11

310

ANGELA

HEANEY

AND JOHN PROCTOR

1200

1000

800L
0

E
E
E600-

400-

200 -

0
J

Time (calendar month)

Figure 2. Mean annual rainfallfor 1 October 1957 - 31 December 1979 forLa Selva Field Station,Costa
Rica. The verticalbars indicate maxima and minima for each month duringthe measurementperiod
(La Selva Field Stationunpublished).

Some aspects of the soils have been described by Marrs et al. (1988). The
parent materials are basaltic and andesitic lavas of Plio-Pleistocene age with a
trend towards tuffor agglomerate-likematerials with increasingelevation. 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 nitrificationshowed 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 junction 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 theirorder

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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 level 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 in Table 1.
RESULTS

The profile diagrams (Figures 3-8) give an impressionof the overall appearance
of the forestsand show the changes in staturewith 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 forestat 10)0m on the VolcainBarvatransect,Costa Rica. Trees

less than 6 m high excluded. Symbols for identifiedtrees over 10 cm dbh: Bi, Brosimumlactescens
(Moore) Berg; Ca, Casearia arborea (Rich.) Urban; Em, Euterpe macrospadixOerst;Ga, Guatteriaaeruginosa Standl.; Gr, G'uarea rhopalocarpa Radlk.; Gx, Guarea sp.; Ho, Hieronymaoblonga var. benthamii
(Tul.) Muell. Arg.; I1,Inga longispicaStandl.; Ip, I. punctata Willd.; Lix, Licania sp.; Mm,Miconia multispicata Naud.; Nn, Naucleopsis naga Pittier;Pa, Pourouma aspera Trecul; Pc, Protiumcostaricense(Rose)
Engler; Pg, P. glabrum (Rose) Engler; Pm, Pentaclethramacroloba (Willd.) Kuntze; Pp, Protiumpanamense (Rose) I. M. Johnston;Qb, Quararibea bracteolosa (Ducke) Cuatr.; Vh, Vochysia hondurensis
Spragne;Wg,WelfiageorgiiWendl.

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312

ANGELA

HEANEY

AND JOHN PROCTOR

40

O-

Figure4. Profilediagram(60 X 7.5 m) of forestat 500 m on the VolcainBarva transect,Costa Rica. Trees
less than 6 m high excluded. Symbols for identifiedtreesover 10 cm dbh: Ax, Ardisia sp.; Cp, Couepia
polyandra (Kunth.) Rose; Cs, Colubrina spinosa Donn. Smith; Dm, Dussia macroprophyllata(D.Sm.)
Harms; Dp, DystovomitapittieriEngl. W.G. D'Arcy; Gy, Guarea sp.; Ig, IriarteagiganteaWendl.;It, Inga
thibaudiana D.C.; Lax, Lauraceae sp.; Mc, MacrolobiumcostaricenseW. Burger;Pac, Parathesischrysophylla Lundell; Rx, Rubiaceae sp.; Ry, Rubiaceae sp.; Vf, VochysiaferrugineaMart.
40

35

W~-

k V

30(-

E~~"'

25 -j~

0A0."

20

15~

0-

Figure 5. Profilediagram(60 X 7.5 m) of forestat 1000 m on the Volcan Barva transect,Costa Rica.
Trees less than 6 m highexcluded. Symbols for identifiedtrees over 10 cm dbh: Bic, Billia colombiana
Planch & Lind.; Ca, Casearia arborea (Rich.) Urban; Ce, Cassipoureaelliptica Poir.; Dp, Dystovomita
pittieriEngl. W.G. D'Arcy; Em, Euterpe macrospadixOerst.;Gg, GuareagrandifoliaDC; 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

~~~~~~~~~~)
C~~~~~~~~~~~~~~~~~~~~~~~~~15-

Figure 6. Profilediagram(60 X 7.5 m) of forestat 1500 m on the VolcainBarva transect,Costa Rica.

Trees less than 6 m highexcluded. Symbolsforidentifiedtreesover 10 cm dbh: Bh, Billia hippocastanum
Peyritsch;Cx, Clethrasp.; Cz, Cyatheaceae; Da, Dendropanax arboreus(L.) Dcne. & Planch.;Dw, Drimys
winteriForst.; Ea, Elaegia auriculata Hemsl.; Ex, Eugenia storkiiStandl.; Go, GuatteriaoliviformisDonn.
Sm.; Hp, Hieronymapoasana Standl.; II, Inga longispica Standl.; Laz, Lauraceae sp.; Mg, Matayba sp.;
Mam, Macrohasseltiamacroterantha(Standl. & L. Wms.) L. Wms.;Ox, Ossaea sp.; Pad, ParathesisadenantheraHook. f.; Pam,Persea americanaMill.; Rf,Rapanea ferruginea(R. & P.) Mez.
30

Figure 7. Profilediagram(60 X 7.5 mn) of forestat 2000 m on the VolcainBarva transect,Costa Rica.
Trees less than 6 m high excluded. Symbols foridentifiedtrees over 10 cm dbh: Ap, Ardisiapalmana
Donn. Smith; Bc, BrunelliacostaricensisStandl.; Cw, Cinclimasp.; Cy, Cyatheaceae;Dip, Didymopanax
pittieriMarch.;Hy, Hamamelidaceae sp.; Hp, Hieronymapoasana Standl.; Mx, Miconia sp.; My,Miconia
occidentalis(SW.) Don; Tx, Tetrorchidiu4m
sp.;
sp.; Mz, Melastomataceaesp.; Px, Piper sp.; To, Tu4rpinia
Vm, Vibu4rnum
mexicanum(nomen).

in differentdiameter classes (Table 3) is remarkable.The frequencyof buttressed treesremains relativelyhigh with increasingplot altitude (Table 4) although
the actual numbersof tall (>1 m high) buttresseson the sampled trees decreases
markedly: there were 22 at 100 m, 39 at 500 m, nine at 1000 m, three at
1500 m, four at 2000 m, and fiveat 2600 m. Lianes decreased in numbers with

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314

ANGELA HEANEY

AND JOHN PROCTOR

30-

i\t2
20- CtA
I~~~~~~~~~~~~~~~~~~~~~I

oL
Figure 8. Profilediagram(60 X 7.5 m) of forestat 2600 m on the Volcin Barva transect,Costa Rica.
Trees less than 6 m high excluded. Symbols for identifiedtrees over 10 cm dbh: Ay, Ardisia sp.; Bc,
Brunellia costaricensisStandl.; Cx, Clethra sp.; Dx, Dendropanax sp.; Iv, Ilex vulcanicola Stand.; Vm,
Viburnummexicanum(nomen).
Table 3. Percentagesof trees(> 10 cm dbh) in a rangeof diameter-classes
on six plots on Volcan Barva,
Costa Rica.
Plot
altitude
(m)

100
500
1000
1500
2000
2600

Diameter-class(cm)
10-20

20-30

30-40

40-50

50-60

60-70

70-80

80-90

90-100

>100

66.7
57.8
59.6
51.4
44.1
51.4

17.0
19.8
18.5
24.5
26.6
22.8

6.2
11.6
8.9

3.1
3.6
6.0
6.5
5.2
7.2

3.1
3.1
2.7

1.8
1.0
1.3
0.9
2.0
2.0

0.6
1.5
1.3

0.2
0.2
1.3

0.2
0.2

0.7
1.0

0.6

0.2
0.3

0.2
1.0
0.2
1.4
1.8
1.6

13.4

16.9
8.5

1.8

2.5
4.6

Table 4. The percentageof sampled trees (N = 100 except for the plot at 2600 m where N = 80) with
buttresses,lianes, skiophyticclimbers,vascularepiphytes,and with more than 50% of the boles covered
by bryophytesat 2 m fromthe ground,in plots at a rangeof altitudeson Volcin Barva,Costa Rica.

Plot
altitude
(m)

100
500
1000
1500
2000
2600

Buttresses
of heights(cm)
50-100

>100

23
23
17
7
14
16

6
8
5
3
3
5

Lianes
of diameter(cm)
>1-5
51
45
33
18
7
14

>5-10

>10

40
25
12
7
4
0

2
7
2
0
0
0

Skiophytic
climbers
75
82
71
95
85
18

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Vascular
epiphytes
89
86
92
99
100
100

Bryophytes
(>50% cover
at 2 m)
5
4
32
79
72
80

Forest on Volca'nBarva

315

140130-

0
0

120-

0
A

110-

100

m
O Om
50
l OOOm
1500m
2 OOOmX
2600m

90*

.0

8070-

560u

cn

50-

403020105

1'5

20

25

Sub-plots (number of 20x20m)

Figure9. Species-area curvesfor trees (> 10 cm dbh) on the six plots on Volcin Barva. (The numbersof
missingspecimensand unidentifiedtrees not included in these curves is givenin Table 1). Each curve
followsthe orderof enumerationof the twenty-five
20 X 20 m sub-plots.

altitude and none >5 cm diameter were recorded fromthe highestplot. Vascular epiphytes were abundant at all altitudes whilst bryophyte cover increased
markedlywith 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 unidentifiedspecimens partiallyaccounts
for the drop in species numbersbetween 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 Mimosaceae 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 in
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 theirpercentageof total basal area and theirnumbersof individualtrees
(> 10 cm dbh) in the six studyplots at a rangeof altitudeson 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
Anacardiaceae
2.15
Annonaceae
1.42
Apocynaceae
0.23
Aquifoliaceae
Araliaceae
3.33
Betulaceae
0.14
Bignoniaceae
Bombacaceae
0.28
Boraginaceae
0.84
Brunelliaceae
Burseraceae
7.29
Caesalpiniaceae 0.99
Capparidaceae
0.05
Caprifoliaceae
Caricaceae
0.05
Celastraceae
Chloranthaceae
Chrysobalanaceae0.04
Clethraceae
Clusiaceae
Combretaceae
Compositae
Coinaceae
Cunoniaceae
Cyatheaceae
Dichapetalaceae
Dilleniaceae
0.45
Elaeocarpaceae
Erythroxylaceae
Euphorbiaceae 4.0
Fabaceae
1.38
Fagaceae
Flacourtiaceae
2.16
Guttiferae
0.16
Hamamelidaceae
Hernandiaceae
1.81
Hippocastanaceae
Humiriaceae
1.25
Icacinaceae
Lacistemataceae 0.05
Lauraceae
1.14
0.58
Lecythidaceae
Malpighiaceae
Malvaceae
0.06
0.04
Margraviaceae
Melastomataceae 0.10
Meliaceae
3.90
Mimosaceae
32.94
Moraceae
5.82
Myristicaceae
1.76
Myrsinaceae
0.14
0.14
Myrtaceae
Nyctaginaceae

4
30
4

3.09
3.82

4
15

0.58

4.80

29

14

3.09

10

0.81

5.40

25

1
4
5

0.24

2
0.35

0.03

4.69

25

1.10

0.26

0.31

8.96

73

47
9
2

4.30
4.06

28
15

1
1

0.43
5.77
0.56

7
17
2

3.63
0.13
0.38
0.64
0.29
3.01
1.22

2
50
3

0.54
0.85
3.98
5.18

6
7
13
17

10
12

0.04
0.04
0.08
0.04
5.40
5.82

1
1
1
1
10
9

11
2

1.82
0.78

11
9

2.92
6.39

2.61
0.05
0.57

3
1
10

1.64
1.18
1.23
0.12

14
2
2
1

1.64
6.92
11.85
3.34
2.23
2.20
0.28

10
32
40
11
5
13
2

5
1
1
11
1
1
2
2
17
69
35
8
3
2

13
2
2
2

0.48

0.03

0.06
3.91

1
16

21.91
25.68
2.32

72
134
4

3.97

15

6.87

65

0.63

10.74

128

0.03

0.12

0.49

3.75
0.16

18
4

1.37
1.23
10.59

3
5
60

0.62

19

12.75
1.74

12
35

0.39

21.74

60

1.68

0.48
6.48

3
24

14.50

63

15
26

2.67
8.49

9
35

5.64
2.65

21
7

3.25

14

2.34

0.06

3.90

14

1.21
0.65
5.82

15
6
19

0.06
4.76

1
22

4.33

32

0.06

0.69

1.12
10.38
10.09
2.66
0.53
0.33
1.03
1.67

13
52
33
21
6
3
24
21

1.10
0.29
6.99
0.08

16
7
35
2

6.83
0.78

61
6

0.28

11

1.70
1.00
0.36

24
18
1

5.10
1.30

39
7

3.24

36

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Forest on Volcan Barva

317

Table 5 - continued

Plot altitudes

Olacaceae
Palmae
Piperaceae
Polygonaceae
Quiinaceae
Rhamnaceae
Rhizophoraceae
Rosaceae
Rubiaceae
Sabiaceae
Sapindaceae
Sapotaceae
Simaroubaceae
Solanaceae
Staphyleaceae
Sterculiaceae
Styracaceae
Symplocaceae
Theaceae
Tiliaceae
Verbenaceae
Vochysiaceae
Winteraceae
Unidentified
or missing

100 m

500 m

1000 m

1500 m

2000 m

2600 m

BA (%) No.

BA (%) No.

BA (%) No.

BA (%) No.

BA (%) No.

BA (%) No.

0.07
0.52

2
11

0.02

0.08

0.11

0.99
10.02

5
114

2.27
0.57

3
7

0.07
0.14
0.12

1
2
2

0.33
0.06
0.40
0.11

1
1
3
2

0.99

11

0.09
2.82
0.94

4
9
2

2.51
0.36
0.71
3.15

50
2
3
8

0.18

0.14

3.66

10

5.10
0.05

3.40
0.04

55
1

0.16

2.88

47

0.05
7.08

1
17

0.70

0.30

2
1
3
1
5

0.09
0.21

11
2

3.27
1.82
3.35
4.66

11.27

10

0.89

10

3.16

18

0.76
3.00
2.07
0.35
3.35

7
30
6
4
10

0.17
2.70
2.87
0.27

1
8
13
2

2.29

6.14

18

0.04
0.49
0.58

1
3
3

2.41
0.11

10
1

0.18

0.52

0.23
14.97

2
54

2.28
7.76

7
32

0.11
4.9

1
36

specimens

DISCUSSION

Forest types
Further informationon 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 classification system of Whitmore (1984) the two
lower plots are probably evergreenlowland rain forest and the other four are
lower montane rain forest although we lack the leaf-size informationwhich is
the most objective criterionfor the classification.
From Hartshorn & Peralta (1987) the plots are said to fall into 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 classificationsystemremainsvague,
however, and objective definitionsin terms of structureand physiognomy are
lacking.
The lowland forests at 100 m and 500 m have a tall stature and 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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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 statement to be reassessed.
It is beyond the scope of this paper to review altitudinal changes in wet
tropical forests in general. Many of the trends in structure and floristicson
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 cauliflory. The most surprising features on Volcan Barva were perhaps that the
largeststatureand most species-richplot was at 500 m ratherthan at the lowest
altitude (100 m) and that the greatestbasal area occurred in the highestplot.
In view of the recent paper on litterfall(Heaney & Proctor 1989) it is useful
here to compare the stature of the Volcain Barva forests with that of other
montane forestswhere production and nutrientcyclinghave 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 summitat 2340 m) (Martin
1977); and Gunung Silam, Sabah (where above 700 m the tallest trees were
21 m) (Proctor et al. 1988). Taller montane forestsin which the trees exceed
the heightof 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 emergentsup 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 unifyinghypothesis which satisfactorilyexplains the
altitudinal changes of the forests on Volcain Barva and the differencesbetween
these forestsand those elsewhere.The likelihood that decreasing nitrogensupply
with altitude is important on Volcain Barva is discussed by Heaney & Proctor
(1989) and Marrset al. (1988). Generalizations fromecological work on Volcan
Barva are hindered because of the non-uniformityof 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-termand 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 currentlybeing monitored for growth
and regenerationstudies by D. and M. Lieberman who are also obtainingmore
detailed climatic informationfromthe mountain.
ACKNOWLEDG

EMENTS

We thankNational 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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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 identifications.Drs P. J. Grubb,
D. L. Kelly and E. V. J. Tanner are thanked for comments on the manuscript.
The work was supported financially by the British Ecological Society, the
Carnegie Tru'st, the Leverhulme Trust and the National Environmental Research Council.
LITERATURE

CITED

ATKIN, L. & PROCTOR, J. 1988. Invertebratesin the litterand soil on Volcan Barva,Costa Rica. Journal of TropicalEcology 4:307-3 10.
BEARD, J. S. 1944. Climaxvegetationin tropical America.Ecology 25:127-158.
BEARD, J. S. 1946. The naturalvegetationof Trinidad. Oxford ForestryMemoir 20, ClarendonPress,
Oxford.
BEARD, J. S. 1949. The natural vegetationof the Windwardand Leeward Islands. Oxford Forestry
Memoir 21, ClarendonPress,Oxford.
BROWN, W. H. 1919. The vegetationof Philippinemountains.Bureau of Science, Manila.
EDWARDS, P. J. & GRUBB, P. J. 1977. Studies of mineralcyclingin a montane rain forestin New
Guinea I. The distributionof organicmatterin the vegetationand soil. Journalof Ecology 65:943969.
FRANKIE, G. W., BAKER, H. G. & OPLER, P. A. 1974. Comparativephenologicalstudies of treesin
tropicalwet and dryforestsin the lowlands of Costa Rica. Journalof Ecology 62:881-919.
GRIEVE, I. C., PROCTOR, J. & COUSINS, S. A. 1990. Soil variationwith altitude on Volcin Barva,
Costa Rica. Catena (in press).
GRIMM, U. & FASSBENDER, H. W. 1981. Ciclos bioquimicos en un ecosistemaforestalde los Andes
Occidentales de Venezuela I. Inventariode las reservasorganicasy minerales(N, P, K, Ca, Mg, Mn,
Fe, Al, Na). Turrialba31:27-37.
GRUBB, P. J. 1977. Controlof forestgrowthand distributionon wet tropicalmountains:with special
referenceto mineralnutrition.Annual Review of Ecology and Systematics8:83-107.
GRUBB, P. J. & STEVENS, P. F. 1985. The forestsof theFatima Basin and Mt Kerigomnaand a review
of montaneand sub-alpineforestselsewherein Papua New Guinea. Departmentof Biogeographyand
Geomorphology,AustralianNational University,
Canberra.
GRUBB, P. J. & TANNER, E. V. J. 1976. The montane forestsand soils of Jamaica: a reassessment.
Journalof theArnoldArboretum57:313-368.
GRUBB, P. J., LLOYD, J. R., PENNINGTON, T. D. & WHITMORE, T. C. 1963. A comparisonof lowland and montane forestin Ecuador. I. The foreststructure,physiognomy,and floristics.Journalof
Ecology 51:567-601.
HARTSHORN, G. S. 1983. Plants. Pp. 118-157 in Janzen, D. G. (ed.). Costa Rican natural history.
Universityof Chicago Press,Chicago.
HARTSHORN, G. S. & PERALTA, R. 1987. Preliminarydescriptionof primaryforestsalong the La
Selva - Volcan Barva altitudinaltransect,Costa Rica. Pp. 281-295 in Almeda, F. & Pringle,C. M.
(eds). Tropical rainforests:diversityand conservation.CalifornianAcademy of Sciences, San Francisco.
HEANEY, A. 1988. Ecological studiesat a rangeof altitudeson VolcdnBarva,Costa Rica. Unpublished
MSc thesis,Universityof Stirling,UK.
HEANEY, A. & PROCTOR, .J. 1989. Chemical elementsin litterin forestson Volcan Barva,Costa Rica.
Pp. 255-271 in Proctor,J. (ed.). Mineralnutrientsin tropicalforestsand savannas.BlackwellScientific
Publications,Oxford.
HOLDRIDGE, L. R., GRENKE, W. C., HATHEWAY, W. H., LIANG, T. & ROSI, J. A. 1971. Forest
environments
in tropicallifezones: a pilot study.PergamonPress,New York.
MARRS, R. H., PROCTOR, J., HEANEY, A. & MOUNTFORD, M. D. 1988. Changesin soils, nitrogen
mineralizationand nitrificationalong an altitudinaltransectin tropical rain forestin Costa Rica.
Journalof Ecology 76:466-482.
MARTIN, P. J. 1977. The altitudinalzonation of forestsalong the west ridgeof GunongMulu. Terminal
reportto Sarawak ForestDepartment,Kuching.
PROCTOR, J., LEE, Y. F., LANGLEY, A. M., MUNRO, W. R. C. & ROBERTSON, F. M. 1988. Ecological studies on Gunung Silam, a small ultrabasicmountainin Sabah I. Environment,foreststructure
and floristics.
Journalof Ecology 76:3 20-340.

This content downloaded on Tue, 5 Feb 2013 02:51:02 AM

All use subject to JSTOR Terms and Conditions

320

ANGELA

HEANEY

AND JOHN PROCTOR

WHITMORE, T. C. 1972. The GunungBenom Expedition 2. An outline descriptionof the forestzones

on northeast GunungBenom.Bulletinof theBritishMuseum (NaturalHistory)D23: 11-15.
WHITMORE, T. C. 1984. Tropical rainforestsof the Far East. (2nd edition). Oxford UniversityPress,
Oxford.
WHITMORE, T. C. & BURNHAM, C. P. 1969. The altitudinalsequence of forestsand soils on granites
near Kuala Lumpur.Malayan NatureJournal22:99-118.

Accepted June 1989

BOOKS
DDT

RECEIVED

and its deriviates - 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.
S. N. & WHITTEN, A. J. 1989. The natural resources, ecology and environKARTIKASARI,
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
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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,
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