Vegetatio 74: 55-80, 1988

© KluwerAcademicPublishers, Dordrecht - Printed in the Netherlands 55

Structure and floristic composition of the lowland rain forest of Los

Tuxtlas, Mexico

E Bongers 1,2, J. Popma 1,2, J. Meave del Castillo 2 & J. Carabias 2

1Permanent address: Department of Plant Ecology, University of Utrecht, Lange Nieuwstraat 106,
3512 P N Utrecht, The Netherlands
2Laboratorio de Ecolog[a, Facultad de Ciencias, Universidad Nacional Aut6noma de Mdxico, 04510
Mdxico DF, Mdxico

Accepted 2.12.1987

Keywords: Allometry, Diversity, Dynamics, Physiognomy, Population structure, Tropical rain forest, Vegeta-
tion structure

Abstract

Physiognomy, structure and floristic composition of one hectare of lowland tropical rain forest was studied
in detail at Los Tuxtlas, Mexico. Physiognomically, the Los Tuxtlas forest should be classified as 'lowland tropi-
cal high evergreen rain forest'. The forest showed a closed canopy at 30-35 m. Of all woody, non-climbing
species with a D B H >_ 1.0 cm 89.407o (94.5070 of all individuals) were evergreen, 25.4070 (59.5 070of the individuals)
had compound leaves, and over 80070 of species (and individuals) had leaves in the notophyll and mesophyll
size classes. The forest structure was characterized by a low density (2976 individuals with a D B H >_ 1.0 cm,
346 individuals with a D B H >_10.0 cm, per ha, excluding vines) with an average basal area (38.1 m 2,
DBH>_ 1.0 cm, 34.9 m 2, DBH>_ 10.0 cm, per ha, excluding vines). This was attributed to the relative maturi-
ty of the forest on the study plot. The study plot contained 234 species (11 208 individuals with a height
>0.5 m), of which 55.1070 (34.8o70 of individuals)were trees, 9.4070 (6.8070)shrubs, 3.4070 (44.3070)palms, 20.1°/0
(5.207o) vines, 6.8070 (8.707o) herbs and 5.10/0 (0.307o) of unknown lifeform. Furthermore, 58 species of epiphytes
and hemi-epiphytes were found. Diversity of trees, shrubs and palms with a D B H _> 1.0 cm was calculated as
Shannon-Wiener index (4.65), Equitability index (0.65), and Simpson index (0.10). The dominance-diversity
curve showed a lognormal form, characteristic for tropical rain forest. The community structure was character-
ized by a relative dominance of Astrocaryum mexicanum in the understorey, Pseudolmedia oxyphyllaria in
the middle storeys, and Nectandra ambigens in the canopy. Species population structures of 31 species showed
three characteristic patterns, differentiated by recruitment: continuously high, discontinuously high, and con-
tinuously low recruitment. Height/diameter and crown cover/diameter diagrams suggested a very gradual shift
from height growth to crown growth during tree development. Forest turnover was calculated as 138 years.
Compared to other tropical rain forests the Los Tuxtlas forest had 1. similar leaf physiognomical characteristics,
2. a lower diversity, 3. a lower density, 4. an average basal area, and 5. a slow canopy turnover.

Nomenclature is given in Appendix I

56

Introduction in a SE to NW direction. It is of volcanic origin, and

In the Neotropics, the northernmost limit in the ge-
ographical distribution of tropical lowland rain for-
est is reached in the state of Veracruz, Mexico. Eco-
logical studies have been conducted in the rain forest
reserve 'Los Tuxtlas' for many years (cf. Gomez-
Pompa & Del Amo 1985; Gomez-Pompa et aL 1976).
An account of the general structure and floristic
composition of the forest is lacking however.
The aim of the present study is to present such an
account, and to evaluate whether the forest at its
northernmost limit is different from other tropical
rain forests, in the neotropics as well as elsewhere.
Study site
The Los Tuxtlas rain forest reserve is situated at the
east side of a mountain range at the coast of the Gulf
o f Mexico in the SE of the state of Veracruz (Fig. 1).
Its elevation varies from 150 m to 530 m. The Los
Tuxtlas mountain range interrupts the coastal plain
dates from the Tertiary up to the Plio-Pleistocene.
The last eruption (of the San Martin Volcano, at
1700 m the highest in the range) was in 1793.
The nearest meteorological station is located in
the village of Coyame located at a distance of 35 km.
At Coyame, the mean annual temperature is 24.6 oC.
The hottest month is May with a mean maximum
temperature of 32.8 °C, and the coldest is January
with a mean minimum temperature of 16.4°C
(Fig. 2). The mean annual precipitation is 4 639 mm.
The distribution of the rainfall over the year is une-
qual. A relatively dry season occurs from March to
May, while a relatively wet season (about 60°70 of the
total annual precipitation) occurs from June to Oc-
tober. The good comparison between the rainfall
curves of Coyame and Los Tuxtlas (Fig. 2) suggests
that the climatic conditions for Coyame can be valid-
ly extrapolated to Los Tuxtlas. The area is affected
by strong and relatively cool winds from the north
which can reach velocities of up to 100 k m / h r and
can cause a drop in temperature down to 10°C for
short periods of time. These winds (known as
'nortes') usually bring heavy rains (up to 3007oof the

..... ~,: Guf of \"~......, Coyame(340) mm

24.6° 4639~800
[281
Montep,~o~ , ~ "~ ~ - -
, uxt,as
Volcano
t /~SanMartrn L~ ~
oC
Santiago~ Sontecomapa~ ~ 40-

Tuxtla I _ ~ - - - - - - ~ y ame 30-

20.
10-
o , ,,o \
km J F M AM J J A S O N D
Fig. 2. Climate diagram o f Coyame following the conventions o f
Fig. 1. Location of the Los Tuxtlas rain forest reserve (95°04 ' to Walter & Lieth (1960-67). The precipitation recorded at the Los
9 5 ° 0 9 ' W and 18°34 ' to 18°36'N). Tuxtlas station is given by the dashed line.
 57

Table 1. Soil characteristics of the study plot (n = 6) a. Guevara-Sada 1985), phenology (Carabias &
Guevara-Sada 1985), seed dispersal (van Dorp 1985)
mean std range
and seed-bank and -rain (Guevara-Sada 1986) had
Organic matter (e/0) 5.49 1.50 3 . 6 8 - 7.50
been carried out there. The study plot was inclined
N (ppm) 5226.5 873.6 3554 - 5766 with the altitude increasing from the SE corner
P (ppm) 2532.7 200.1 2265-2851 (140 m) to the NW corner (165 m). Each of the 100
K (ppm) 926.4 131.4 7 2 8 . 7 - 1122.0 10 × 10 m subplots into which the study plot was
pH 6.27 0.16 6.1-6.5
divided had an inclination of at least 10%. In addi-
Sand (e/0) 75.7 7.0 66 - 86
Silt (%) 15.3 7.8 6 - 26
tion to this, two small runnels originate in the plot,
Clay (%) 9.0 3.5 6 - 16 causing very steep slopes locally. These runnels
usually dry up during the dry season.
a Soil samples were taken at six sites within the plot. At every site Within each subplot all individuals with a di-
five subsamples (15 cm depth, excluding litter layer) were taken
ameter at breast height ( D B H ) > _ 1.0 cm were
with a 8 cm diameter soil sampler. These were mixed carefully
to form the sample used for analysis. The samples were dried at
marked with an aluminium tag. Because palms were
room temperature. Organic matter was determined according to very abundant in the understorey these were tagged
the method of Walkley-Black. pH was measured at a 1:2.5 and measured separately when they did not have a
soil/water ratio. Total N, P and K were determined using a modi- stem at 1.3 m but did have one at 0.5 m height. Of
fied Kjeidahl method. Salicic acid was added to bind nitrates,
every tagged individual species name, lifeform and
and NaESO 4 was used to raise the boiling temperature of the di-
gestion. Ion concentrations were measured in an autoanalyzer.
diameter at breast height ( D B H ) were recorded. For
The analysis of soil texture was done using the hydrometer meth- all non-climbing plants also height (H), lower crown
od. limit ( L C L ) , largest crown diameter (D1) and the di-
ameter perpendicular to D 1 (D2) were measured.
yearly total), and are frequent from December to Height measurements were done with a marked pole
February. up to 5 m, and with a Suunto clinometer for higher
The substrate consists mainly of igneous rocks trees. Crown cover was calculated according to the
(basalt and andesite), mixed with volcanic ash. The formula of an ellipse,
soils derived from this material show poorly devel-
oped profiles, and contain high amounts of organic C = 0.25 • D l • D 2 , ,.x (1)
material (Chizon 1984). They are classified as vitric
andosols (Tv 20-1bc, FAO/UNESCO 1975). The crown volume as an ellipsoid,
soil at the study site is neutral to weakly acid, and,
according to the standard of reference given by V = 0.167. D 1 • D 2 • ( H - L C L ) . 7r (2)
Young (1976) very rich in N, P and K (Table 1). Or-
ganic matter content is 5.5%. Its textural composi- All individuals which reached a height of 0.5 m and
tion can be classified as a sandy loam. did not have a D B H >_ 1.0 cm, were identified and
counted per quadrat. Epiphytes were recorded on an
abundance scale in the whole (100 x 100 m) study
Methods plot: less than 5 individuals, rare (R); 5 - 1 0 , occa-
sional (O); 11-30, frequent (F); 31-100, abundant
Field work was carried out from March to November (A); and over 100 individuals, very abundant (VA).
1983, with additional short periods during 1984 to As far as possible individuals were identified to
1986. A study plot of 1 ha, containing all major species level. All identifications were checked a sec-
regeneration phases (e.g. gap, building phase and ond time in the field. Species that could not be iden-
mature forest) was chosen in a forest section without tified were assigned to morphospecies. O f all mor-
signs of recent human disturbance. This particular phospecies sterile speciments were collected, and
plot was of special interest because several other stored in the collection of J. Meave del Castillo. In
studies, notably on productivity (Alvarez & the text, species and morphospecies will be used as
58
separate taxa except where stated otherwise.
For all species ( D B H _> 1.0 cm) leaf size was deter-
mined measuring 20 leaves (leaflets in the case of
compound leaves) taken randomly from one mature
individual. The pinnate leaves of the palms found in
Los Tuxtlas are considered here as compound. Leaf
sizes were grouped into Raunkiaer's leaf size classes
as modified by Webb (1959). Species were considered
to be deciduous when mature individuals remained
leafless for a longer period (i.e. 2 to 14 weeks around
the beginning of the dry season). Different degrees
of deciduousness were not distinguished.
Species richness and three indices o f diversity were
calculated, the Shannon-Wiener index H, the Simp-
son index C and the Equitability index E:
H = - ~ Pi 2LOG Pi (3)
where Pi = ni/N; /'/i = number o f individuals of
species i, and N = total number of individuals.
C = ~, pi 2 (4)
E = H/H m (5)
where H m = 2LOG S; S = number of species.
A standard method for analysing dominance-
cliversity structure is to plot the logarithm of some
importance value o f the species on the ordinate
against their rank order (number in the species se-
quence from most to least important) on the abscissa
(Whittaker 1975). A problem is to find a good meas-
ure of importance.'The measures commonly used are
density, frequency, biomass, basal area, cover, or
combinations of these. The use of combinations is
considered an advantage, as various parameters of
the species' importance are taken into account.
However, a high combination value does not distin-
guish between a high value for one parameter and
a high value for another one: both can result in the
same high combination value. To avoid this problem
to some extent two parameters were used here
separately to calculate the species rank order: a dis-
tribution index (calculated as density, #/ha, times
frequency, % of 10 × 10 m plots), and cover sum.
Frequency diagrams of species with at least
15 individuals with a DBH_> 1.0 cm (or trunk
height _>0.5 m for palms) were constructed. Size-
class limits and number o f classes were different per
species. The number of classes was determined as
M = INT (5.1°LOG N) (6)
where Mis the number of classes, and Nis the popu-
lation size. D B H class width was then obtained by
dividing the D B H range by the number of classes,
including the upper limits in each class.
Allometric relations between D B H and height of
individuals were described using an empirical model
suggested by Kira (1978, see also Yamakura et al.
1986),
Hma x • D B H
Height - DBH + K (7)
where Hmax is a size constant (asymptotic maxi-
mum height), and K is a shape constant ( D B H at
which half of the asymptotical maximum height is
reached), defining the curve. The relation between
D B H and Cover was described using a cubic poly-
nomial regression model. Regressions were calculat-
ed on mean values f o r D B H i n t e r v a l s , to reduce com-
putational requirements and effects of outlying
values.
From 1983 to 1986 all new gaps were recorded and
mapped. In 1985 gaps were counted and gap sizes
were measured, using the vertical projection of the
canopy opening as a measure of gap size (Brokaw
1982). In January 1986 the percentage of the total
plot area in gap phase (Pg), building phase (Pb) and
mature forest (Pro) was determined according to the
method o f Martinez-Ramos et al. (in press). Turno-
ver rates were calculated following Hartshorn
(1978). Based on recent gap creation data, and as-
suming that gap phase survives ca. 1 yr, the turnover
rate R was calculated as:
Rg = lO0/Pg (8)
Assuming that the building phase survives for 25 to
35 yr (Martinez-Ramos 1985), turnover rate may
als,3 be calculated as:
25 × lO0/(Pg + Pb) < Rub <
35 × lO0/(Pg + PO) (9)
 59

Results (25.407o of the species) had compound leaves (Ta-

ble 2). The latter figure was high, due to the domi-
Physiognomy and vegetation structure nance of understorey palms. When only large in-
dividuals are taken into account (DBH >_30 cm, no
The closed canopy of the Los Tuxtlas rain forest palms reach this limit), deciduousness increased to
reaches a height of 30-35 m, with occasional in- 15.0°70 of the species, while the percentage with com-
dividuals up to 40 m. Real emergent trees are absent, pound leaved species increased to 32.5o7o of the spe-
also outside the study plot. The 2976 individuals of cies. Eleven of the 15 deciduous species were canopy
trees, palms, and shrubs (DBH_> 1 cm) had a sum species. The proportion of deciduous species and of
cover of 365o7o of the plot area (Table 2), which me- compound-leaved species thus increased towards the
ans that above an average point in the study plot be- upper canopy. In all leaf size spectra between 81 and
tween three and four crowns were located. This fig- 93O7o fell into the combined noto- and mesophyll
ure might have been even higher if epiphytes and class (Table 2). When using different bases for the
especially lianas, which can have large crowns, calculation of spectra, the results can vary consider-
would have been included. The basal area was ably.
38.1 m E which corresponds with 0.38o70 of the plot
area (Table 2).
Of all individuals with a DBIt>_ 1.0 cm, 5.5°7o FIoristics and diversity
(10.6°70 of the species) were deciduous, while 59.5°7o
A total of 292 species of vascular plants were found
in the plot (Table 3). This figure excludes some herb
Table 2. Percentage of deciduousness, compoundness, and leaf species that did not reach a height of 0.5 m. In total,
size spectra weighted by species, abundance, basal area, crown 72 families and 181 genera could be identified. The
cover and crown volume a.
tree group comprised most species, and in abun-
# # Basal Crown Crown
dance it was only surpassed by the palms (Table 4).
Species Indiv. area cover volume Among trees the most important families in terms
(m 2) (m 2) (m 3) of number of species were Leguminosae
(22 species), Moraceae (11), Rubiaceae (10) and
Deciduous 10.6 5.5 15.5 10.3 14.3
Lauraceae (8). There was a high number of woody
Compound 25.4 59.5 23.0 39.4 29.4
liana species (Table4), mainly Bignoniaceae
(9 species), Malpighiaceae (5) and Sapindaceae (5).
Size class b A few small stature palm species were very abundant
Nanophyllous 2.1 0.2 0.4 0.3 0.2
Microphyllous 8.5 1.8 7.5 5.2 6.5
Notophyllous 38.7 22.3 56.1 47.3 57.2 Table 3. S u m m a r y of the floristic survey of the study plot a.
Mesophyllous 42.3 70.0 29.1 42.4 31.3
Macrophyllous 7.8 5.6 6.3 4.6 4.7 Identified 070 of °70 of
Megaphyllous 0.7 0.1 0.7 0.2 0.2 to level of morpho- individuals
species
Total (10007o) 142 2976 38.07 36466 181728
Species 72.2 97.5
aOnly trees, shrubs and palms with a D B H >_ 1 cm; values are Genus 9.4 1.0
percentages Family 9.4 0.8
bRaunkiaer's size classes as modified by Webb (1959). Unknown 9.8 0.7
Nanophyllous = 0.25 - 2.25 cm 2,
microphyllous = 2.25 - 20.25 cm 2, N (100070) 234 11208
notophyllous = 20.25 - 45.0 cm 2,
mesophyllous = 4 5 . 0 - 182.25 cm 2, aIncluding individuals over 0.5 m high, excluding epiphytes and
macrophyllous = 1 8 2 . 2 5 - 1640.25 cm 2, hemiepiphytes; the latter belong to 58 additional taxa (56 species
megaphyllous > 1640.25 cm 2. identified to species level, 2 identified to genus level).
60

Table 4. Lifeform spectrum for the study plot 1. to decide whether they belonged to the shrubs, trees
or vines group. Vascular epiphytes were relatively
DBH>_ 1 cm Height_>0.5 m
scarce (M.A. Soto-Arenas, pers. comm.) and com-
Lifeform 2 % of % of % of °/0 of
prised mainly species of Orchidaceae (19 species),
species indiv, species indiv. Araceae (12), Bromeliaceae (5) and Cactaceae (4). A
complete list of species and their density in the plot
? 0 0 5.1 0.3 is given in Appendix I.
H 0 0 6.8 8.7
Species richness and some diversity measures were
V 23.2 11.0 20.1 5.2
P 2.7 48.1 3.4 44.3
compared for the same plot, using different plant
S 9.7 5.0 9.4 6.8 and plot-size limits and excluding or including
T 64.3 35.9 55.1 34.8 different lifeforms (Table 5). It is clear that depend-
ing on the survey characteristics, values for exactly
N (100%) 185 3344 234 11208
the same site can show a very large variation.
IEpiphytes and hemi-epiphytes (58 species) excluded because
absolute density was not determined.
2?: unknown, H: herbs, V: vines, P: palms, S: shrubs, T: trees Dominance - diversity relation
of various sizes.

The differences in species rank using the distribution

in the understorey, especially Astrocaryum mexica-
num. The palms group comprised fewest species, but
was the largest in terms of abundance (Table 4). The
herb species Dieffenbachia seguine, Spathiphyllum
cochlearispathum, Aphelandra aurantiaca, and
several fern species, were found on the forest floor.
Common shrub species were Schaueria calycobrac-
tea, Acalypha skutchii and Myriocarpa longipes,
which may be called small gap pioneers (cf. Popma
et al., 1988a). The group of unknown lifeforms
was formed by woody species of which only
juveniles were found, in which case it was impossible
index or the cover index is shown in Fig. 3. Tentative-
ly, three groups of species can be distinguished:
1. Species with a similar distribution rank as their
cover rank are found along the diagonal in the
figure (difference between both ranks not higher
than 50%, an arbitrary chosen limit, species
marked by a cross in Fig. 3). These are mainly
sub-canopy to understorey species which seem to
regenerate successfully in small gaps or even in
full shade.
2. Species with a high cover (low cover rank) but
relatively poorly distributed, thus showing a high
distribution rank (difference in ranks greater

Table 5. Diversity measures and survey characteristics I.

Lifeforms included Size limit Area (ha) N S H E C

T, V, S, P, H 0.5 cm height 1.0 11208 234 5.12 0.65 0.074

T, V, S, P 1 cm DBH 1.0 3344 185 5.07 0.67 0.081
T, V, S, P 1 cm DBH 0.5 1584 136 4.97 0.70 0.082
T, S, P 1 cm DBH 1.0 2976 142 4.65 0.65 0.101
T 1 cm DBH 1.0 1202 119 5.68 0.82 0.034
T, V, S 10 cm DBH 1.0 359 88 5.31 0.82 0.054
T, V, S 10 cm DBH 0.5 176 58 4.97 0.85 0.058
T 20 cm DBH 1.0 180 52 4.59 0.80 0.086
T 40 cm DBH 1.0 75 31 3.95 0.80 0.137

tLifeforms: T = trees, V = vines, P = palms, S = shrubs, H = herbs

N = number of individuals, S = number of species, H = Shannon-Wiener index, E = Equitability index, C = Simpson index.
 61

than 50%, species marked by a square in Fig. 3). In Fig. 4 the hierarchy of species based on their cover
These species are mainly large canopy or sub- is shown. The distribution index curve (not shown)
canopy trees of low abundance that seem to was similar to the cover curve.
regenerate only in larger gaps.
. Species which are well distributed (low distribu-
tion rank) but with a relatively low cover, and thus Allometry
a high cover rank (again, difference in ranks ex-
ceeding 50%, species marked by a triangle in Figure 5a shows the models fitted to the D B H -
Fig. 3). These are mainly small understorey spe- height and DBH-cover relation for all trees in the Los
cies that show good regeneration in the full shade. Tuxtlas one hectare plot. Both models showed a
good fit (p < 0.001, R 2 > 0.98). The increasing scat-
130- ter around the regression lines with increasing D B H ,
a oo o~ +++4-~ + +~+ + +4444++++~+
120- can be attributed to the low number of trees in the
110- larger size classes due to the limited plot size. An in-
100- 23 24
crease in D B H is at first coupled to an increase in
oo oo o o + + + -~- 4+ + + + +++÷ +
90- tree height, with a more or less constant, small cover.
80-
+ +
Later on, the pattern is completely reversed: when
1810~O20021 022 + + + + + + +

70-
the trees reach the canopy layer (above 25 m) stem
17
[] + ++ + + +
+
+
++
++ diameter increase is coupled to an increase in cover,
60-
+ +
+ ÷ + +
zx
with almost no further increase in height. Three pat-
¢6 50-
16
u
+ + +
+
£x
terns for individual species, representing different
40- ~s + + + +
+ + z~ ecological groups, are given as well (Fig. 5). In gener-
30- lo ~.~ ~,
+12 al, the species curves do not deviate very much from
1

+ 34
the total community pattern. The shade tolerant un-
7 +3
1 o .k -
~03 +5 6 30 32
3~ derstorey species Sapranthus microcarpus (An-
Oi
0
,
10 20
,
30
i ,
40
,
50 60
i
70
,
80
i ,
90 100110120130140
, , i i , nonaceae) has thicker stems and larger crowns at a
Species rank (cover) certain height than the total community pattern
would predict. When individuals approach maturi-
Fig. 3. Species rank according to the distribution index value (y-
ty, growth in diameter is favoured compared to
axis) vs. species rank according to the cover value (x-axis). Trian-
gels indicate species for which cover rank is at least twice their
growth in height. Individuals of understorey species
distribution index rank. 25: Chamaedorea tepejilote, like S. microcarpus are mature at a smaller height
26: Sapranthus microcarpus, 27: Trophis mexicana, 28: Bactris than individuals of canopy species, hence the thicker
trichophylla, 29: Chamaedorea schiedeana, 30: Orthion oblan- stems. The subcanopy species Pseudolmedia oxy-
ceolatum, 31: Pouteria spec., 32: Aegiphila costaricensis,
phyllaria (Moraceae) and the canopy species Nec-
33: Acalypha skutchii, 34: Capparis bacluca, 35: Myriocarpa
Iongipes. tandra ambigens (Lauraceae) have thinner stems and
Squares indicate species for which distribution index rank is at smaller crowns at a certain height, as compared to
least twicetheir coverrank. 13: Nectandraambigens, 14: Guarea the total community. Nectandra ambigens
glabra, 15: Spondias radlkoferi, 16: Trichilia martiana, (Lauraceae) has relatively smaller crowns at a certain
17: Dialium guianense, 18: Vatairealundellii, 19: Ficus colubri- trunk diameter.
nae, 20: Coccoloba barbadensis, 21: Lonchocarpus guatemalen-
sis, 22: Cordia stellifera, 23: Ficus insipida, 24: Dussia mex-
icana.
Crosses are species with comparable ranks for both parameters. Species population structure
l: Astrocaryum mexicanum, 2: Pseudolmedia oxyphyllaria,
3:Poulsenia armata, 4: Dendropanax arboreus,
The population structures of sufficiently abundant
5:Cymbopetalum baillonii, 6: Faramea occidentalis,
7:Mortoniodendron guatemalense, 8: Psychotria simiarum, species (31 of 142 species) were analysed. Three
9:Croton schiedeanus, 10: Turpinia oceidentalis, 11: Pouteria general patterns, suggesting different population dy-
durlandii, 12: Rheedia edulis. namics in the forest, could be distinguished. The
62

+_1
t3A3
+_2

1000- 23~+~4'-24
[3181917

1 0 ~ 9 22
" '~'~ ~ 3o
' i ~ * +1'1-'/"~33
~ 35
£kl 100-
E

>
o z~'++4"~++4+~4,~-.N.+4,,~_+%++.1.+
o 10-
4+~z,,,,~.+
-H-

-14-

0.1
1'0 2'0 3'0 4'0 5'0 6'0 7'0 8'0 90 160 li0 1;~0 130 140
species rank

Fig. 4. Dominance - diversitycurve. Speciesordered from high to low cover (x-axis) vs. their cover (y-axis). Symbolsand numbers as
in Fig. 3.

first group (Pattern type I in Fig. 6a) is formed by
all species having the highest frequency in the first
or second DBHclass, showing a gradual decrease in
the number of individuals towards the bigger classes.
Eleven species (35.5%) showed this pattern. Fig-
ure 6b shows the population structure of Pseudo#
media oxyphyllaria as an example of a type I curve.
In the second group (Pattern type II in Fig. 6a) the
first class has the highest frequency value, but the ad-
jacent classes (second and/or third) are badly
represented. Frequency rises again more or less sh.ar-
ply in intermediate classes and declines at the upper
limit of the DBHrange. Eight species (25.8%) pres-
ent this pattern. As an example Fig. 6c shows the
pattern of Cymbopetalum baillonii. The third group
(Pattern type III in Fig. 6a) is the largest group,
formed by 12 species (38.7o/0) and all big canopy spe-
cies abundant enough to be included in this analysis
belong to it. A type III population curve is charac-
terized by the concentration of the greater part of the
individuals (in general 500/o or more) in the first size
class, while the remaining classes are poorly and very
uniformly represented (reversed 'J' curve). Extreme
examples are Poulsenia armata (Fig. 6d) and Mor-
toniodendron guatemalense, two typical high cano-
py species which had respectively 90.7% and 88.5%
of their individuals situated in the first D B H class.
Nectandra ambigens, the most abundant species in
the canopy of the forest, was included in this group
in spite of its apparently different histogram. This
species shows a tendency of a continuous increase
towards the bigger DBHclasses, having the smallest
classes badly represented. However, this species
forms a huge seedling bank with a high mortality
(C6rdova 1985). Only very few individuals survive
and reach the size needed to be included in our anal-
ysis. Probably, most other canopy species present the
same type III pattern.
 63

mean height (m) Total community . m e a n c o v e r (m2) height (m) cover(m2)

Pseudolmedia oxyphyllaria
(a) 800
36- (c)
25.0-
++~ +
33-
22.5-
30- 900 ÷ + +
200- + ++ ++ + 600
27-
175- ++ + +
24-
÷ + ÷ +
21- 15.0-
600
18- ÷ 4- + + + 400
+# / 12.5-
15-
12-
300
10.0

7.5
q .
9-
200
6- 5.0- ++ + 4-

3-
25'
0
10 2 0 3 0 4 0 5 0 6 0 7 0 8 0 90 100 110 1 2 0 1 3 0 1 4 0 0.0
D B H (cm)
0 ,5 I0 I'5 2'0 2'5 30 3'5 40
DBH Ccm)

height [m) Sapranthus m i c r o c a r p u s c o v e r (m2) height (m) cover (m2)

Nectandra ambigens
13-
(b) -400 35.0-
(d)
+
12- ~+
32.5. * ++ +
+ +
11- 30.0. ÷ 900
10- -300 27.5
+ + + +++ ++
9-
8-
7-
++
+ +
+
+ +
++
25.0.
225.
20.0-
+ +++: !+.÷+ 600
+ + "200
6- + + 17.5-
+*+++ + + +
5- + + 150 °
4- + -~++ + + + 125-
-100 10.0.
3- 300
2- ,~J+ 7.5-
1- 5.0-
0" 25- Xx - ~ x xx ,
0
0.0
D B H (cm)
g 1}3 1'5 2'0 2'5 3'0 3'5 4'0 4'5 5'0 5'5 60 6'5 7'0
D B H (cm)

Fig. 5. Allometrix relations: height and cover versus diameter (a) Total c o m m u n i t y patterns for D B H versus Height ( - - ) and D B H versus
Cover ( . . . . . . ). Points are m e a n values for D B H intervals, lines are regressions (Height = (43.3 x D B H ) / ( D B H + 33.9), p < 0.001,
R 2 = 0.96, Cover = 1.803 x D B H + 0.0059 x D B H 2 + 0.00017 x DBI-P, p < 0.001, R 2 = 0.97); (b) Scatter diagram for Sapranthus
microcarpus (all individuals). Lines are regressions for the total community, except S. microcarpus (Height =
(43.6 x D B H ) / ( D B H + 34.1), p < 0.001, R 2 = 0.96, Cover = 1.80 x D B H + 0.0060 x D B H 2 + 0.00017 × DBI-P, p < 0.001, R 2 = 0.95);
(c) Scatter diagram for Pseudolmedia oxyphyllaria (all individuals). Lines are regressions for the total c o m m u n i t y , except R oxyphyllaria
(Height = (44.3 x D B H ) / ( D B H + 36.2), p < 0.001, R 2 = 0.97, Cover = 1.65 x D B H + 0.0086 x D B H z + 0.00016 x DBI-P, p < 0.001,
R 2 = 0.96); (d) Scatter diagram for Nectandra ambigens (all individuals). Lines are regressions for the total c o m m u n i t y , except N. ambi-
gens (Height = (45.4 x D B H ) / ( D B H + 37.2), p < 0.001, R 2 = 0.95, Cover = 1.76 x D B H + 0.0128 x D B H 2 + 0.00012 x D B H 3,
p < 0.001, R 2 = 0.95).

Gap dynamics 4 yr o f age was estimated to be 2.9% (a mean o f

0.725% annually), the total area o f building phase
In January 1985 the plot contained 11 small canopy as 13.6%, and the area o f mature forest as 83.5%.
gaps, o f which 7 were between 5 and 10 m 2, one was Canopy turnover rates o f the plot are 138 y r ( R g ,
ca. 25 m 2, two were ca. 50 m 2 and one was ca. based o n recent gaps), and between 152 and 212 yr
75 m 2. In January 1986 the total area o f gaps up to (Rgb, based on area in gaps and building phase).
64

% A %
type [
44
hypothetical curve types 30 Pseudolmedia oxyphyllaria
(N=134)

20

1I 10.
....... .... ......................
.
13
6 1
I I I
4.7 8.4 12,2 15,9 19.623,427.130.834.638.342.C

DBH upper limit of DBH-class (cm)

100
% % D
type ][ C1 90 .m type Ill
~ Cymbopetalum baillonii 80- Poulsenia armata
30- (N=86)
70-
60-
20- 50-
40-
30-
10-
20
10-
1 1 1 1 1 2 1
5.3 9.5 13.8 18.0 22.3 26,5 30.8 35.0
0 8.2 15.4 22.6 29.837.0 44.2 51.4 58.665.873.£

upper limit of DBH-class (cm) upper limit of DBH-class (cm)

Fig. 6. Three types of population structure. Type I, Good reproduction and continuous recruitment; Type II, Good reproduction and
discontinuous recruitment; Type III, Good reproduction but bad recruitment. A. Hypothetical curve types. B. Population structure of
Pseudolmedia oxyphyllaria, Type I. Other species in this group are: Chamaedorea tepefilote, Astrocaryum mexicanum, Bactris
trichophylla, Aegiphila costaricensis, Acalypha skutchii, Piper hispidum, Trophis mexicana, Psychotria chiapensis, Myriocarpa lon-
gipes, and Capparis baduca. C. Cymbopetalum baillonii, Type If. Other speciesin this group are: Pouteria spec., Sapranthus microcarpus,
Faramea occidentalis, Piper lapathifolium, Psychotria simiarum, Guarea glabra, Quararibea guatemalteca and Chamaedorea schiedea-
nus. D. Poulsenia armata, Type II1. Other species in this group are: Mortoniodendron guatemalense, Dendropanax arboreus, Turpinia
occidentalis, Croton schiedeanus, Pouteria durlandii, Psychotria flava, Psychotria faxlucens, Rheedia edulis, Amphitecna tuxtlensis,
Orthion oblanceolatum and Nectrandra ambigens.

Discussion studies, one is forced to recalculate m a n y

The description o f tropical rain forest vegetation
and subsequent c o m p a r i s o n with other forests
presents several limitations. It is very difficult to
directly c o m p a r e results o f studies o n tropical rain
forests, because o f the heterogeneity in criteria and
m e t h o d s used. W h e n c o m p a r i n g results o f different
parameters. This is a very laborious and in m a n y
cases even an impossible task. Plot sizes used are
mostly smaller than 1 ha (circa 60% o f the studies
mentioned in A p p e n d i x II). W h e n larger plots are
used the lower plant size limit is mostly 10 cm D B H
(cf. A p p e n d i x II). It m a y be doubted whether plots
smaller than 1 ha can give a representative picture o f

the community in a tropical rain forest vegetation.
On the scales used, populations of a large majority
of the species present are very small. Also, the sheer
impossibility of including lianas and epiphytes can
cause severe bias in quantitative descriptions of for-
est structure.
Gap dynamics
Forest turnover rate is regarded as an important de-
terminant of forest structure and floristic composi-
tion (Bazzaz 1984; Whitmore 1984; Brokaw 1985;
Newbery et al. 1986) and is therefore discussed first.
In forests with a short turnover time (high distur-
bance rate) one might expect 1. higher densities of
pioneer species (species which complete their entire
life cycle in gaps), 2. higher plant density, and
3. individuals to be of smaller size on average than
in forests with a long turnover time, other conditions
being the same. The gaps found in our plot were
small. Canopy closure will thus occur mainly by
(lateral) growth of existing individuals, not by
colonizing pioneer species (Bazzaz 1984; Martinez-
Ramos & Alvarez-Buylla 1986). The proportion of
mature forest in the study plot is high, compared to
nearby plots analysed by Martlnez-Ramos et al. (in
press). They found that, over a 3 year period in a
5 ha plot, on average 1.61% of the canopy is opened
up by treefall annually, and that at any moment 42%
of the area was covered by gap or forest in the build-
ing phase. This results in an Rg of 62 yr, and an Rg b
between 60 and 83 yr. These values overlap, so that
their plot can be regarded as constant in its internal
(relatively high) disturbance rate for the last 25 to
35 yr. Our plot, on the other hand, shows a n Rg of
138 yr and a n Rgb between 152 and 212 yr. This
could indicate a recent increase in disturbance in our
plot, although it is still less disturbed than the plot
of Martinez-Ramos et al. (in press). A reason for this
might be that our plot is situated on a somewhat less
steep slope, and it is relatively protected from the
strong northern winds because of its aspect. The
turnover time Rg calculated for out plot, 138 yr, is
relatively high in comparison to values reported for
other Central American tropical rain forests (range
62 to 155 yr; Brokaw 1985). This suggests that the
65
study plot is situated in a relatively undisturbed part
of the forest. The difference between our results and
those of Martinez-Ramos et al. (in press) for nearby
plots (distance 200-700 m) indicates that even on
this small scale large differences in gap dynamics ex-
ist. This may lead to considerable differences in
many structural and other characteristics of the for-
est.
Physiognomy
Community physiognomy is generally described by
means of spectra of characteristics such as leaf size,
compound versus simple leaves, evergreen versus
deciduous habit, bark morphology, presence versus
absence of buttresses, etc. Most spectra given in the
literature are based on the number of species and do
not weight for the actual quantity of any such
characteristic in the community. Weighting by struc-
tural parameters of the vegetation, like abundance,
basal area, crown cover and crown volume, give a
more realistic approximation, each emphasizing
different aspects of the community structure.
The Los Tuxtlas forest can be classified as high
tropical lowland evergreen rain forest (cf. Miranda
& Hernandez-X 1963). The adjective evergreen indi-
cates that the forest as a whole has an evergreen ap-
pearance, not that all species are evergreen. The
deciduous species present are mostly canopy species.
Subcanopy and understorey species are nearly al-
ways evergreen. The difference between evergreen
and semi-evergreen forests depends on the percent-
age of deciduous species in the forest. No general
rule exists. Gomez-Pompa (1973) mentions that at
least 80% of the species in the 'high evergreen selva'
is evergreen. Hall & Swaine (1976, 1981) find, based
on abundance, an evergreen percentage of more than
80 for their evergreen forest types in Ghana (only
taking into account mature individuals of
megaphanerophytes). When compared to other
tropical rain forests, the Los Tuxtlas forest has a high
percentage of evergreenness (Table 6). Its percentage
of compound species is a little lower than in some
other evergreen rain forests (Table 6), although
differences are rather small. Often, compond leaves
and deciduousness are seen as adaptations to sea-
66

Table 6. Proportion of deciduous and compound-leaved spe- sonal d r o u g h t (cf. G i v n i s h 1978). I f so, the latter
cies: comparison with other lowland tropical rain forests a. does n o t seem to be a n i m p o r t a n t e n v i r o n m e n t a l fac-
t o r in Los Tuxtlas- the m a j o r i t y o f the species o f Los
Location Size limits o7ocompound 07odeciduous
Tuxtlas, 99 (69.7o70), are simple-leaved a n d ever-
species (indiv.) species (indiv.)
green. O f the r e m a i n i n g species, 28 (19.7°70) are
Mexicoe dbh -> 1 cm 25.4 (59.5) 10.6 (5.5) compound-evergreen, 7 (4.9o/0) are simple-
dbh -> l0 cm 26.9 (18.6) 16.7 (10.1) deciduous, a n d 8 (5.6°70) are c o m p o u n d - d e c i d u o u s .
h ->6 m 29.2 (32.3) 14.6 (7.9) Givnish (1978) s u p p o s e d t h a t c o m p o u n d leaves
c o u l d be a d a p t a t i v e when r a p i d vertical g r o w t h
Costa Ricaf upper story - 27
lower story - 7.5 a n d / o r high t u r n o v e r o f leaves a n d b r a n c h e s is need-
total - 17 ed (e.g. early successional species colonizing light
gaps). This hypothesis is n o t s u p p o r t e d by the results
Panamag 20b o f this s t u d y however, as o f the 20 p i o n e e r species
in the plot, o n l y 4 have c o m p o u n d leaves.
Trinidadh emergents 47.6 (43.7) 26.2 (3.4)c
canopy 30.5 (53.3) 8.7 (1.3) l e a f size spectra for Los Tuxtlas are c o m p a r e d to
lower story 17.6 (31.2) 8.0 (0.8) t h o s e for o t h e r t r o p i c a l l o w l a n d rain forests in Ta-
ble 7. W i t h the exception o f Shinguipino, E c u a d o r
Ecuadori h_>6 m 36 ( G r u b b e t al. 1963), all forests s t u d i e d have m o r e
t h a n 75°70 o f the species in the n o t o p h y l l plus
BrazilJ dbh _>10 cm 30.7
m e s o p h y l l size classes. The Los Tuxtlas rainforest is
Brazilk dbh -> 10 cm 28.8d thus very similar to the o t h e r rain forests, as far as
its l e a f size spectra are c o n c e r n e d . In general, leaf
Ghana l size in t r o p i c a l rain forests is negatively related to
plot a dbh ->10 cm - (19)
l a t i t u d e a n d altitude, a n d positvely to a n n u a l rainfall
plot b (22)
(Webb 1968; D o l p h & Dilcher 1980). F u r t h e r m o r e ,
Philippines m upper story 32 large-leaved species seem to be favoured u n d e r an
middle story 17 e q u a l d i s t r i b u t i o n o f rainfall t h r o u g h o u t the year, a
lower story 13 high n u t r i e n t avilability, a n d a m o d e r a t e r a d i a t i o n
New Guinean dbh ->10cm 21, 16, 29, 27 intensity (Givnish 1984).

aValues are percentages of species, values in parenthesis are per-

centages of individuals
bExcept Ficus ssp
clncluding semi-deciduous species
dof all mega, meso, micro and nano phanerophyte species
eThis study, Los Tuxtlas, lowland rain forest
fFrankie et al. (1974), La Selva, lowland rain forest
gCroat (1978), Barro Colorado Island, tropical forest
hBeard (1946), Trinidad, evergreen seasonal forest, excluding
the Mora type. Emergents: _>100 ft, canopy: 40-90 ft, lower
storey: 10- 30 ft (only individuals with a DBH -> 10 cm)
iGrubb et al. (1963), lowland tropical forest
JCain & de Oliveira Castro (1959), but study plot from Pires et
aL (1953), Tierra firme tropical rain forest
kCain et a1.(1956), Tierra firme tropical rain forest
1Hall & Swaine (1976), (a) west evergreen rain forest (b) moist
evergeen rain forest
mBrown (1919), in Richards (1952), tropical dipterocarp rain
forest
npaljmans (1970), tropical rain forest, plot 1-4.
Structure
Figures 7 a n d 8 show d e n s i t y a n d b a s a l area for vari-
ous l o w l a n d t r o p i c a l rain forests, c o m p a r e d with the
Los Tuxtlas value for exactly the s a m e p l a n t size
limits. G e n e r a l i n f o r m a t i o n o n the studies used for
c o m p a r i s o n is given in A p p e n d i x II. C o m p a r e d with
m o s t o t h e r forests, either in A m e r i c a , A f r i c a o r
A s i a , the Los Tuxtlas forest has a low density. F o r
densities o f larger trees ( D B H >_ 20 cm), Los Tuxtlas
values are similar to those r e p o r t e d f r o m elsewhere.
C o m p a r i s o n o f Figs. 7 a n d 8 shows that, in general,
a high vegetation density c o r r e s p o n d s with a high
b a s a l area, b u t the r e l a t i o n s h i p is weak (i.e. for
D B H > _ 10 cm" R = 0.581, P < 0.01). This m e a n s
t h a t a high d e n s i t y can be c o m b i n e d with a.high bas-
 67

Table 7. Leaf sizes: comparison with other lowland tropical rain forests a.

Location Size limits Leaf size classes

Leptophyllous Nanophyllous Microphyllous Notophyllous Mesophyllous Macrophyllous Megaphyllous

Mexico ~ dbh >- 1 cm 0 (0) 2.1 (0.2) 8.5 (1.8) 38.7 (22.3) 42.3 (70.0) 7.8 (5.7) 0.7 (0.1)
dbh >_ 10 cm 0 (0) 2.6 (0.6) 9.0 (5.0) 46.2 (55.6) 33.3 (33.1) 7.7 (5.0) 1.3 (0.6)
h >_6 m 0 (0) 3.1 (0.6) 9.4 (3.8) 41.7 (47.6) 38.5 (43.4) 6.3 (4.3) 1.0 (0.3)

Trinidad e Emergents 0 2.4 (0.0) 4.8 (20.5) 85.7 (83.5) 7.0 (16.0) 0
Canopy 2 (34) 0 17.4 (8.5) 76.1 (55.2) 4.3 (1.9) 0
lower story 2.2 (34.3) 0 11.8 (16.4) 80.4 (71.5) 7.8 (12.1) 0

Ecuador f h >_6 m - 0 9 14 50 27 0

Brazilg dbh >_ 10 cm 2.0 2.0 13.3 74.0 7.3 1.3

Brazil h dbh >_ 10 cm 2.9 0.7 12.3 75.5 8.6 0

Brazil i dbh >_ 10 cm 1.2 (0.5) 0 (0) 14.7 (10.6) 78.5 (86.0) 5.1 (2.7) 0.6 (0.7)

NigeriaJ dbh >_ 10 cm 0 0 10 84 6 0

Philippines k ?0 0 4 86 10 0
New Guinea I dbh>_ 10 cmb 3.5, 7, 2, 1 27.0, 29.0 69.5, 64, 76, 79.5
22.0, 12.5

Australia m c

type 1 0 0- 5 3 0 - 50 5 0 - 70 0 0
type 2 0 30 40 30 0 0

a Values give percentage of species, values in parenthesis give percentage of individuals.

b Microphyllous is smaller or equal to microphyllous, mesophyllous is larger or equal to mesophyllous; microphyllous/notophyllous
and notophyllous/mesophyllous groups are splitted up in two parts: half in each neighbouring class.
c Ignoring shrubs, palms, and deciduous species.
d This study, Los Tuxtlas, lowland rain forest.
e Beard (1946), Trinidad, evergreen seasonal forest, excluding the Mora type. Emergents: >- 100 ft, canopy: 4 0 - 9 0 ft, lower story:
10 - 30 ft (only individuals with a DBH >- 10 cm).
f Grubb et aL (1963), lowland tropical forest.
s Cain & de Oliveira Castro (1959), but study plot from Pires et al. (1953), Tierra firme tropical rain forest.
h Cain et al. (1956), Tierra firme tropical rain forest.
i Mori et al. (1983), tropical moist forest.
J Richards (1952), wet evergreen forest.
k Brown (1919), in Richards (1952), tropical dipterocarp rain forest.
J Paijmans (1970), tropical rain forest, plot 1 - 4 .
m Webb (1959), 1, Mesophyll vine forest, 2, semi evergreen vine forest.

al area (Meave del Castillo 1983; Holdridge et al.
1971; Lescure et al. 1982; Mori et al. 1983), but also
with an intermediate or even low one (Hall & Swaine
1981). On the other hand, some of the forests studied
by Sarukh~tn (1968) show a low density but a high
basal area due to a relatively high number of very
thick canopy trees. The Los Tuxtlas forest has an in-
termediate basal area (see Fig. 8). Rollet (1978) gives
pantropical means for tropical lowland evergreen
rain forest of 21 m2/ha (DBH>>_ 10cm) and
7 m2/ha ( D B H > _ 6 0 cm). Compared to these
values, Los Tuxtlas has a higher basal area (34.9 m 2,
68

y=x A
/
/
I /
/
c-- / r'-
400- O
32 a 24 22185/231:~,5~ 9 15 28 13f19 10 ~
i , 'i ' ' ' I
o l, i,
300" /
O /
._1 7 22"27// O16
15 -r-
200" ~10~D'22~9 1~ ~'~8 ~13 20
25
I 12 29 ,-.,,,~ 3o ~
t- lOOi 35~
40

0 L/ / 2 7
0 100 2(}0 36O 46O 56O 66O 760 860 960
Density for other forests (ha -1)

4000, /
B
/
/ y=X
/
t- /
15 / 11
3000. o/ / 0 O -1 t-
/ o
X /
I-- /
12/
O- .2 ~
2000- 7 /
13// 13 2.5 m
a:::l
1
[]
"3.3
[] [] zn/.q~_ 2 .~.o ~
1000- 12,)°"16 o? -'4.5-
GO / -5
C
O> /
/
/
/ I
l obo 20bo 30'00 4000 50'00 6000
Density for other forests (ha -1)

Fig. 7. Comparison of Los Tuxtlas vegetation density with other lowland tropical rain forests. Density for Los Tuxtlas (y-axis) using the
same plant size limits as used for the reported value (x-axis). Numbers refer to A p p e n d i x I I . I n case values for several forest types are
reported the ranges are shown by a connecting line. Symbols: v = A s i a , A = A f r i c a , o = South America, [] = Central America. A :
lower D B H limits _> 10 c m , B: l o w e r D B H limits < 10 c m .

D B H >_ 10 cm, and 16.5 m 2, D B H >_ 60 cm, per ha) lowland forests of America, Africa or Asia, com-
due to a relatively high number of thick trees. The pared to Los Tuxtlas, neither a tendency that differ-
location of the study plot in a rather undisturbed ences are related to differences in latitude can be
part of the forest, may partly explain the low densi- found. This is in agreement with Gentry (1982), who
ty/intermediate basal area values. The variation found that density does not differ significantly be-
among studies is very high. There seem to be no con- tween tropical sites.
sistent differences in density or basal area between The vertical structure of the Los Ttlxtlas rain forest
 69

//y= x
/
/
40- /
I
'4
13
~n 3~/ 02
11
,¢ ~.3
-q. ~4..0
04 821 20 28 31 6 14 930 / 4 19 33 5 4.5
E m ~ 7R~ go oF 9 o t~ 10
t~
13 016 I1! / -15
/011 -20
x /
30- 26
V /
/
09 / -30
0 /
d /
s /
/ I'-
O
/
O)
/
i,,_ /
/ ~D
/ O3
20- I
09
123
3

20 3'0 40 5'0 6'0 74.9

Basal area for other forests (m2.ha -1)

Fig. 8. Comparison of Los Tuxtlas basal area with other lowland tropical rain forests. Basal area for Los Tuxtlas (y-axis) using the same
plant size limits as used for the reported value (x-axis). Numbers refer to Appendix II. In case values for several forest types are reported
the ranges are shown by a connecting line. Symbols: v = Asia, a = Africa, o = South America, ta = Central America.

will be described elsewhere (Popma et al. 1988b). Rico, and by Richards (1939) in Nigeria. The plots
studied by Lescure et al. (1982), Prance et al. (1976)
and two of the four plots studied by Proctor et al.
Diversity (1983), are much richer in species. Very few studies
report values for the three diversity indices (Table 8).
The large variation in plot sizes and plant size limits It seems that in Los Tuxtlas H a n d C are higher, while
causes serious problems when comparing diversity E is lower in comparison to other rain forests, sug-
among forests, because most diversity measures are gesting a lower overall diversity. Several explanations
highly dependent on those sizes and size limits (Ta- can be given for this relatively low diversity. The
ble 5, Letouzey 1978; Rollet 1978). Conseqeuntly, study plot is situated in a relatively undisturbed part
the only way to compare reported values realistically of the forest, hence the low proportion of pioneer
is to use exactly the same plot size and plant size limit species (14.1% of the species, 6.6o7oof the individuals
for each comparison made. Because in Los Tuxtlas with a DBH_> 1 cm). This is in accordance with
one hectare was studied in detail, comparisons could what Connell (1978) postulated: the forests richest
be made with studies using other plant - and plot in species should be those with an intermediate dis-
size limits too, as long as they concern data for plots turbance regime, because they contain both pioneer
equal or smaller than one hectare and D B H _> 1 cm and shade tolerant species. Various observations on
or plant height _>0.5 m. The values reported in the temperate as well as on tropical forests support this
literature are compared with the Los Tuxtlas values intermediate disturbance hypothesis, although
for exactly the same plot and plant size limits. The others do not (Whitmore 1982). Also, soil nutrient
Los Tuxtlas forest is relatively species-poor (Fig. 9), concentrations are high (Table 1). According to
except in comparison to the forests studied by Saru- Huston (1979; 1980), both regular disturbance and
kh~m (1968) in Mexico, by Crow (1980) in Puerto low nutrient availability promote high diversity.
70

Others found a positive correlation between species taker 1975). Plotting the symbols of Fig. 3 in Fig. 4
diversity and soil fertility (Richards 1952; Whitmore clearly shows which species are over- or under-valued
1984) including phosphorus (Gartlan et al. 1986), using the cover rank order in comparison to the dis-
and according to Ashton (1977) species richness tribution rank order. The combination of the two
reaches a maximum on soils with intermediate fertil- figures gives a clear picture of the relative impor-
ity. Species richness has been shown to have a posi- tances of the species in the community as well as of
tive correlation with precipitation in forests studied the role certain species types play in determining
by Hall & Swaine (1981) and Holdridge et al. 1971). community structure. Three species have a relatively
Gentry (1982) found that this was only the case when high cover. These species reach a different maximum
taking all individuals with a D B H >_2.5 cm into ac- height. Astrocaryum mexicanum is an understorey
count, and not when this limit was increased to species (8 m), Pseudolmedia oxyphyllaria is a sub-
10 cm DBH. Another reason might be that the spe- canopy species (24 m), and Nectandra ambigens is
cies pool is limited: many neotropical rain forest spe- a canopy species. When mature, individuals of these
cies may not reach the area in their geographical dis- species exclude each other spatially. The same ef-
tribution. fects, although less pronounced, occur in the re-
The dominance-diversity curve (Fig. 4) seems to maining part of the curve.
conform to a lognormal species distribution (rela-
tively few species with a high or a low importance,
many species with intermediate ones). This kind of Species populations
curves is characteristic for species-rich communities
including tropical rain forest (Hubbell 1979; Whit- Because estimates of tree ages are difficult to obtain

/
/ / y =X
69
/
/
x /
/
i.- /
/

d
8, oo- /
/ 16a
0
/
/
S 22 32
01 1
28
if) 0 18 0 0
7
/ ,,~ ,r-
1
// 2 33
/ D V30a
O
//o .o 16b
1 / 9b
o 50-
/ v
/ 30b
.IQ /
/
E /
Z ,4 5

/
/
/
0/"
0 5b 16o 26o
Number of species for other forests

Fig. 9. Comparison of Los Tuxtlas species richness with other lowland tropical rain forests. The x-axis gives the species richness for other
forests as reported in the literature. On the y-axis the species richness for Los Tuxtlas calculated for the same size and area limits as used
in the study to which it is compared. Numbers refer to Appendix II. In case values for several forest types are reported the ranges are
shown by a connecting line. Symbols: v = Asia, A = Africa, o = South America, o = Central America. Areas of plots: studies nr. 4,
5, 7:0.1 ha; nr. 1:0.2 ha; nr. 2, 11:0.25 ha; nr. 30:0.32 ha; nr. 9B: 0.5 ha; nr. 3:0.72 ha; nr. 33:0.8 ha; nr. 8, 9A, 14, 16, 17, 18, 19, 22,
28, 32:1.0 ha. Lower plant size limits (DBH): study nr. 11:1.0 cm; nr. 7:2.5 cm; nr. 1, 2:3.3 cm; nr. 3:4.0 cm; nr. 30A: 4.5 cm; nr. 16A:
5.0 cm; nr. 4, 5, 8, 9A, 9B, 14, 17, 18, 22, 28, 30B, 32, 33:10.0 cm; nr. 16B: 15.0 cm.
 71

Table 8. Diversity components: comparison with other lowland rain forestsa

Location Area b DBI-I b S H E C

b L. Tuxtlasc b L. Tuxtlasc b L. Tuxtlasc b L. Tuxtlasc

Mexico d 0.25 3.3 78.3 61.3 4.76 3.65 0.76 0.62 - -

Puerto Rico e 0.72 4.0 51 90.0 4.0 3.62 0.70 0.56 - -
Costa Rico f ? 2.5 57 43.2 5.40 - 0.046 -
Venezuelag Plot A 0.5 10.0 79 54.5 5.4 4.89 0.86 0.85 0.04 0.063
Venezuelag Plot B 0.5 10.0 63 54.5 5.3 4.89 0.89 0.85 0.04 0.063
Venezuelag Plot C 1.0 10.0 83 79 4.8 5.17 0.75 0.82 0.08 0.058
Brazil h 1.0 10.0 108 79 6.23 5.17 0.92 0.82 0.031 0.058
Brazil i 1.0 10.0 87 79 5.37 5.17 0.83 0.82 0.044 0.058

a S = number of species, H = Shannon-Wiener index, E = Equitability index, C = Simpson index.

b area (ha) for which values are given, and lower D B H limit (cm).
c Los Tuxtlas value for the same area and D B H limit, where possible the mean value for various plots of that plot size/plant size
is given.
d Meave del Castillo (1983). Values are means for four plots.
e Crow (1980). Values calculated from diversity/evenness curves.
f Knight (1975). Values are means for five old forest plots of various sizes. S value for 0.1 ha, H and C values given but not
comparable to Los Tuxtlas because exact area is not given.
g Uhl & Murphy (1981).
h Pires et al. (1953).
i Black et al. (1950).

in regions with more or less continuous growth, an
adequate description of tree growth can only be
made using long term increment data (see Lieber-
man & Lieberman 1985, and references there). How-
ever, using allometric relations between parameters
that are known to be correlated with age like DBH,
tree height and cover, it is possible to analyse the way
in which trees grow in the forest.
Hall6 et al. (1978) use double logarithmic plots of
height versus diameter, where a line H = 100 x D
( H = h e i g h t in m, D = stem diameter in cm)
separates the so-called model conforming trees
(trees of the future) found at the left hand side of the
line, from strongly reiterated trees (trees of the pres-
ent) found at the right hand side. Their division
seems arbitrary however, and no justification for the
position of the dividing line is given. Torquebiau
(1981) describes other lines to distinguish the differ-
ent sets of trees, and states that the position of the
line probably depends on the particular forest stud-
ied. When comparing Kira's (1978) height/diameter
plots, it becomes clear that the form of the curves
is highly variable. Inspection of other curves (Schulz
1960; Takeuchi 1961) shows the same phenomenon,
indicating that there is no justification for simple
rules of thumb.
When compared to the graphs presented by Kira
(1978), the Los Tuxtlas forest is lower than most
others, in spite of its much higher precipitation.
Asian rain forests are generally higher than neotropi-
cal rain forests (Whitmore 1984), but maybe the low-
er stature of the Los Tuxtlas forest can partly be ex-
plained by the absence of emergent trees. Possibly
tree dimensions are limited by the strong and cold
northern winds that occur frequently during the
winter period.
The graphs from the total community as well as
from the individual species suggest a gradual shift
in allocation of resources during tree development,
rather than discrete passages from one stage of their
development to another. Whitmore (1984, p. 20)
states that 'height growth of most tree species is
complete when only between one-third to one-half
of the final bole diameter has been reached'. Our
results do not support this conclusion, as tree height
increases monotonically with D B H (Fig. 5).
72
For the description of the size structure of popula-
tions of different species it is not very accurate to use
uniform diameter classes for the construction of fre-
quency diagrams. In most studies the same size-class
limits are used for small sized species as for large
sized ones. This results in a differentiation between
large and small stature species rather than between
species with a different population structure (e.g.
Knight 1975). Consequently it is better to construct
a separate diagram for each species, taking into ac-
count the population size and the maximum D B H
obtained in the plot. Following this procedure, three
different types of population structure were distin-
guished (Fig. 6a). The 11 species with a type I popu-
lation structure (34.4% of all studied species) are
small or medium sized trees. These species appear to
have good reproduction and regular recruitment in
the study plot, and their density decreases rather
regularly with increasing size (Fig. 6b). The type II
group contains species which show good reproduc-
tion but discontinuous recruitment into larger size
classes (Fig. 6c). Three possible explanations may be
mentioned. Firstly, some size classes could be more
critical than others having a higher size specific mor-
tality. Second, growth rates may be size-dependent,
causing the peculiar down-up-down shaped histo-
grams of this group. Thirdly, these species may de-
pend on irregularly occurring events for their
regeneration. The species with a type III population
curve are considered to have a good reproduction but
a poor recruitment into large DBHclasses (Fig. 6d).
This groups includes many seedling-bank forming
canopy species.
Knight (1975) provides an analysis of population
structures in rain forest in Panama. Comparability
to our data is limited due to the fact that Knight
(1975) joined data of many separate plots, not taking
into account that the population structure of one
species can be different at different sites. Also,
Knight (1975) used the same size class limits for the
construction of the histograms for all species. Con-
sequently, he distinguishes a pattern corresponding
to small sized species, not taking into account the
size structure of their populations. In the present
study small stature species were dispersed over the
three types discussed before. There are two addition-
al groups described by Knight that were not found
in the present study: one of them having the highest
frequency in intermediate classes, and the other one,
his 'unfrequent reproducers', having the better
representation in the bigger DBHclasses. These two
groups could be artefacts produced by the exclusion
of seedlings and small plants from Knight's (1975)
analysis. Probably these patterns can be considered
as Type II or Type III curves respectively. The same
holds for the seven population structure patterns
given by Rollet (1978). Including smaller size classes
in Rollet's (1978) analysis would probably cause
most of his curve types to merge with either of the
three patterns described here.
Acknowledgements
The authors are indebted to G. Ibarra and S. Sinaca,
without whose help identification of many species
would have been impossible. S. Castillo provided
valuable assistance in the field. M. A. Soto-Arenas
censused and identified the epiphytes. We thank the
staff of the Los Tuxtlas Biological Station for their
continuous support. The IBM Scientific Center of
Mexico and the Department of Experimental Plant
Ecology of the Catholic University of Nijmegen,
The Netherlands, kindly provided computer facili-
ties. We thank E. van der Maarel and M. J. A. Werg-
er for their encouragement, and for critically revis-
ing the manuscript. F. Bongers and J. Popma were
supported by grant nr. W84-204 of the Netherlands
Foundation for the Advancement of Tropical
Research (WOTRO) to M. J. A. Werger, Utrecht and
E. van der Maarel, Uppsala.
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A P P E N D I X I. SPECIES LIST
Complete list o f plant species and morphospecies found on one hectare of tropical evergreen lowland rain forest at Los Tuxtlas, state of
Veracruz, Mexico. Species sorted alphabetically, by family. 23 morphospecies (79 individuals) could not be identified, not even on the
family level. Speciments of unknown or partially unknown species are being kept in the collection of J. Meave del Castillo. The species
were identified either in the MEXU herbarium of the Institute o f Biology, Universidad Nacional Aut6noma de M6xico, or in the herbarium
of the Los Tuxtlas field station, or by G. Ibarra-Manrlquez. Nomenclature of tree species is in agreement with Ibarra-Manrlquez (1985).
Epiphytes were identified by M.A. Soto-Arenas.
Data following species name are Lifeform (T = tree, V = vine, S = shrub, P = palm, H = herb, E = epiphyte, EH = hemi-epiphyte)
and abundance (all individuals over 0.5 m in height included, values between brackets indicate the number of individuals with a DBH
greater or equal to 10 cm). In the case of the epiphytes and hemi-epiphytes, only an estimate of the abundance is given (R = rare, < 5
ind.; O = occassional, 5 - 1 0 ind.; F = frequent, 11-30 ind.; A = abundant, 31-100 ind.; VA = very abundant, >100 ind.).

ACANTHACEAE Anthurium scandens (Aubl.) Engl. E F

Acanthaceae spec. 02 H 3 Anthurium schlechtendalii Kunth E A
Aphelandra aurantiaca (Scheidw.) Lundell H 76 Dieffenbachia seguine (L.) Schott H 108
Odontonema callistachyum (Schlecht. & Monstera acuminata C. Koch EH V
Cham.) Kuntze S 23 Monstera tuberculata Lundell EH O
Schaueria calycobractea Hilsenbeck & Philodendron guttiferum Kunth E F
Marshall S 292 Philodendron inaequilaterum Liebm. EH V
Philodendron radiatum Schott E F
ANACARDIACEAE Philodendron srnithii Engl. E F
SpondiasradlkoferiDonn. Sm. T 14 (5) Philodendron tripartiturn (Jacq.) Schott EH R
Spathiphyllum cochlearispathum (Liebm.)
ANNONACEAE Engl. H 228
Annonaceae spec. 01 T 3 (1) Syngonium macrophyllum Engl. EH F
Cymbopetalum baillonii R.E. Fr. T 58 (22) Syngonium popophyllum Schott EH F
Malmea depressa (Baill.) R.E. Fr. T 1
Sapranthus microcarpus (Donn. Sm.) R.E. ARALIACEAE
Fr. T 195 (4) Dendropanax arboreus (L.) Decne. &
Planch T 57 (21)
APOCYNACEAE Oreopanax obtusifolius (L.) L.O. Wms. EG O
Aspidosperma megalocarpon Muell. Arg. T 14 (2)
Forsteronia viridescens Blake V 73 (1) ASTERACEAE
Sternmadenia donnell-smithii (Rose) Wood- Asteraceae spec. 01 V 1
son Y 12 (7) Eupatorium galeottii Robins T 2
Eupatorium spec. 01 V 4 (1)
ARACEAE Liabum discolor Benth. & Hook. v 2 (1)
Anthurium flexile Schott ssp. flexile EH A Neurolaena lobata (L.) R. Br. S 7
76

Zexmenia suberosa Villasenor CHRYSOBALANACEAE

Couepia polyandra (HBK.) Rose T 1
BIGNONIACEAE
Amphilobium paniculatum (L.) HBK. var. CLUSIACEAE
paniculatum V 1 Clusia spec. 01 E R
Amphitecna tuxtlensis A. Gentry T 41
Anemopaegma chrysanthum Dugand V 7 COMBRETACEAE
Bignoniaceae spec. 04 V 7 Combretum laxum Jacq. V 4
Bignoniaceae spec. 05 V 1
Bignoniaceae spec. 06 V 3 COMMELINACEAE
Bignoniaceae spec. 09 V 1 Campelia zanonia (L.) HBK. H 19
Callichlamys latifolia (L.C. Rich.) K.
Schum. V 34 (2) CONNARACEAE
Paragonia pyramidata (L.C. Rich.) Bur. V 2 Connarus schultesii Stand. ex R. Schultes V 43
Stizophyllum riparium (HBK.) Sandwith V 5
CONVOLVULACEAE
BLECHNACEAE Ipomoea spec. 01 V 5
Lomariopsis recurvata Fee EH F Ipomoea phillomega (Vull.) House V 3

BOMBACACEAE DICHAPETALLACEAE
Bernoullia flammea Oliv. T 1 (1) Dichapetallum donnell-smithii Engl. V 8 (1)
Quararibea funebris (Llave) Vischer T 14 (2)
Quararibea guatemalteca (Donn. Sm.) DILLENACEAE
Standl. T 44 (1) Davilla aspera (Aubl.) Ben. var. matudae
(Lundell) L.O. Wms. V 31
BORAGINACEAE
Cordia megalantha Blake T 2 (1) ERYTHROXYLACEAE
Cordia stellifera I.M. Johnson T 3 (3) Erythroxylon tabascense Britton 1 (1)

BROMELIACEAE EUPHORBIACEAE
Aechmea bracteata (Sw.) Griseb. E F Acalypha diversifolia Jacq. S 5
Aechmea nudicaulis (L.) Griseb. E A Acalypha skutchii I.M. Johnst. S 207
Catopsis cf. sessiliflora (Ruiz & Pavon) Croton schiedeanus Schlecht. T 52 (8)
Mez E R Omphalea oleifera Hemsl. T 5 (1)
Tillandsia dasyliriifolia Baker E R Sapium nitidum (Monach.) Lundell T 2 (1)
Vriesia gladioliflora (Wendl.) Ant. Wiener E O
FLACOURTIACEAE
BURSERACEAE Casearis nitida Jacq. T 2
Bursera simaruba (L.) Sarg. 3 (2) Lunania mexicana Brandegee T 7 (1)
Pleuranthodendron lindenff (Turcz.) Sleu-
CACTACEAE mer T 32 (6)
Disocactus mmulosus (Salm-Dyck) Kim-
nach E R GESNERIACEAE
Epiphyllum oxypetalum (DC.) Haw. E O Columnea purpusii Standl. E V
Epiphyllum pumilum (Vaupel) Britt. & R. E F
Rhipsalis baccifera (Mill.) W.T. Stearn E V GUTTIFERAE
Calophyllum brasiliense Camb. T 1
CAPPARIDACEAE Rheedia edulis (Seem.) Triana & Planch. T 27 (4)
Capparis baduca L. T 46 (1)
Crataeva tapia L. T 19 (2) HIPPOCRATEACEAE
Salacia megistophylla Standl. V 136
CELASTRACEAE
Celastraceae spec. 01 S 1 ICACINACEAE
Crossopetalum parviflorum Hemsl. T 5 Mappia longipes Lundell T 2 (1)
Rhacoma spec. 01 T 5
 77

LAURACEAE MARCGRAVIACEAE
Lauraceae spec. 01 T 1 (1) Souroubea exauriculata Delpino E R
Lauraceae spec. 02 T 1 (1)
Lauraceae spec. 04 T 5 MELASTOMATACEAE
Licaria spec. 01 T 1 Clidemia setosa (Triana) Gleason H 4
Nectandra ambigens (Blake) C.K. Allen T 76 (42) Miconia sp. nov. S 2
Nectandra spec. 01 T 1 (1)
Ocotea dendrodaphne Mez. T 12 (4) MELIACEAE
Ocotea spec. 01 T 1 Guarea bijuga C. DC. T 38 (2)
Guarea glabra Vahl. T 29 (13)
LEGUMINOSAE Guarea grandifolia A. DC. T 19 (4)
Acacia mayana Lundell T 1 Triehilia martiana C. DC. T 9 (5)
Albizia purpusii B. & R. T 3 (1) Trichilia pallida Sw. T 60
Cynometra retusa B. & R. T 22 (2)
Dialium guianense (Aubl.) Sandw. T 4 (3) MENISPERMACEAE
Dussia mexicana (Standl.) Harms. T 860 (2) Abuta panamensis (Standl.) Krukoff & Bar-
Erythrina folkersii Krukoff & Moldenke T 2 neby V 19 (1)
Inga brevipedicellata Harms T 3 Menispermaceae spec, 01 T 1
Inga jinicuil Schlecht. T 11
Inga quaternata Poepp. T 4 MONIMIACEAE
Inga sapindoides Willd. T 9 Mollinedia viridiflora Tul. T 12
1riga spec. nova T 12 (1) Siparuna andina (Tul.) A. DC. S 4
Inga spec. 01 T 6
Inga spec. 03 T 1 MORACEAE
Inga spec. 04 T 7 Brosimum alicastrum Sw. spp. alicastrum T 25 (2)
Leguminosae spec. 01 T 1 Cecropia obtusifolia Bertol. T 4 (2)
Leguminosae spec. 02 T 1 Clarisa biflora Ruiz & Pavon ssp mexicana
Lonchocarpus cruentus Lundell T 1 (1) (Liebm.) W. Burger T 4 (2)
Lonchocarpus guatemalensis Benth. T 4 (3) Ficus aft. cotinifolia HBK. T 1 (1)
Machaerium cobanense Donn. Sm. V 1 Ficus colubrinae Standl. T 3 (3)
Machaerium floribundum Benth. V 21 (4) Ficus insipida Willd. T 2 (2)
Pithecellobium arboreum (L.) Urban T 3 (1) Ficus obtusifolia HBK. E F
Pterocarpus rohrii Vahl. T 3 (2) Poulsenia armata (Miq.) Standl. T 275 (8)
Swartzia guatemalensis (Donn. Sin.) Pittier T 8 Pseudolmedia oxyphyllaria Donn. Sm. T 443 (57)
Vatairea lundellii (Standl.) Killip T 4 (3) Trophis mexicana (Liebm.) Bur. T 254 (5)
Trophis racernosa (L.) Urb. T 2
LOGANIACEAE
Strychnos tabascana Sprange & Sandwith V 4 MUSACEAE
Heliconia spec. 01 H 25
LYCOPODIACEAE
Lycopodium cf. dichotomum Jacq. E R MYRSINACEAE
Myrsinaceae spec. 01 S 3
MALPIGHIACEAE Oerstedianthus brevipes (Lundell) Lundell S 37
Bunchosia lindeniana Adr. Juss T 43 Parathesis lenticellata Lundell S 7
Heteropteris laurifolia (L.) Adr. Juss. V 2 (1)
Hiraea fagifolia (DC.) Adr. Juss. V 12 MYRTACEAE
Malpighiaceae spec. 01 V 40 Eugenia spec. 01 T 7
Malpighiaceae spec. 02 T 6 Eugenia spec. 02 T 3 (1)
Malpighiaceae spec. 03 V 3 Eugenia spec. 03 T 14 (1)
Malpighiaceae spec. 08 V 1 Eugenia spec. 04 T 16

MALVACEAE NYCTAGINACEAE
Hampea nutricia Fryxell T 29 Neea psychotrioides Donn. Sm. T 8 (1)
Robinsonella mirandae Gomez-Pompa T 3 (1) Pisonia aculeata L. V 6

MARATTIACEAE OCHNACEAE
Danaea nodosa (L.) Smith H 188 Ouratea tuerckheimii J.D. Smith T 41
78

ORCHIDACEAE Diplazium lonchophyllum Kuntze H 220

Chysis bractescens Lindl. E R Niphidium crassifolium (L.) Lellinger E R
Dichaea muricata (Sw.) Lindl. E R Pleopeltis revoluta (Spreng. ex Willd.) A.R.
Encyclia cochleata (L.) Lemee E O Smith E A
Epidendrum imatophyllum Lindl. E R Polypodium spec. 01 E O
Epidendrum isomerum Schltr. E R Polypodiaceae spec. 01 H 1
Epidendrum polyanthum Lindl. E O Polypodiaceae spec. 02 H 19
Gongora aff. quinquinervis Ruiz & Pavon E R Polypodiaceae spec. 03 H 3
Lycaste cochleata Lindl. ex Paxt. E O Polypodiaceae spec. 04 H 32
Maxillaria pulchra (Schltr.) L.O. W m s E R
Maxillaria tenuifolia Lindl. E O RHAMNACEAE
Nidemia boothii (Lindl.) Schltr. E V Gouania lupuloides (L.) U r b a n V 2
Oncidium luridum Lindl. E R
Ornitocephalus inflexus Lindl. E R RUBIACEAE
Pleurothallus lewisae Ames E R Faramea occidentalis (L.) A. Rich. T 71 (8)
Pleurothallus microphylla A. Rich. & Gal. E V Hamelia longipes Standl. T 3
Restrepiella ophiocephala (Lindl.) Garay & Hillia tetrandra Sw. E F
Dunsterv. E A Psychotria chagrensis Standl. T 8 (1)
Scaphyglottis dubia (A. Rich & Gal.) Psychotria chiapensis Standl. T 25
Benth. & Hook. ex Hems E A Psychotria faxlucens Lorence & Dwyer T 65 (2)
Sobralia decora Bateman E O Psychotria flava Oerst. & Standl. T 29
Trigonodium egertonianum Batem. ex Psychotria limonensis Krause S 7
Lindl. E F Psychotria papantlensis (Oerst.) Hemsl. T 14
Psychotria simiarum Standl. T 41 (8)
PALMAE Psychotria veracruzensis Lorence & Dwyer T 14
Astrocaryum mexicanum Liebm. P 2324 Randia aft. laetervirens Standl. V 10
Bactris trichophylla Burret P 220 Randia pterocarpa Lorence & Dwyer T 26
Chamaedorea ernesti-augustii H. Wendl. P 7 Randia retroflexa Lorence & Nee V 15
Chamaedorea schiedeana Mart. P 1011
Chamaedorea tepejilote Liebm. P 1079 RUTACEAE
Desmoncus aft. ferox Bartlett P 3 Zanthoxyllum kellermanii P. Wilson T 1 (1)
Geonoma oxicarpa Mart. P 7
Reinhardtia gracilis (H. Wendl.) Burret P 310 SAPINDACEAE
Allophylus campostachis Radlk. T 24 (5)
PASSIFLORACEAE Cupania dentata DC. T 4
Passiflora spec. 01 V 1 Cupania macrophylla A. Rich. T 5
Paullinia costaricensis Radlk. V 2 (1)
PIPERACEAE Paullinia pinnata L. V 34
Peperomia nigropunctata Miquel E V Sapindaceae spec. 01 V 1
Peperomia petenensis Trelease E V Sapindaceae spec. 02 V 2
Peperomia quadrifolia (L.) HBK. E V Sapindus saponaria L. T 2 (2)
Peperomia rotundifolia (L.) HBK. E V Thinouia tomocarpa Standl. V 9
Piper aduncum L. S 5
Piper aequale Vahl S 18 SAPOTACEAE
Piper aff. amalago L. S 3 Dipholis minutiflora Standl. T 3 (3)
Piper amalago L. T 13 (2) Pouteria campechiana (HBK.) Baehni T 7 (2)
Piper hispidum Sw. S 41 Pouteria durlandii (Standl.) Baehni T 38 (4)
Piper lapathifolium Steud. S 38 Pouteria sapota (Jacq.) H.E. Moore &
Piper nitidum Vahl S 5 Stearn T 4
Potomorphe umbellata (L.) Miq. S 1 Pouteria spec. 01 T 110 (1)
Sapotaceae spec. 01 T 5
POLYGONACEAE
Coccoloba barbadensis Jacq. T 3 (1) SMILACACEAE
Smilaxspec. O1 V 1
POLYPODIACEAE
Campyloneurum angustifolium (Sw.) Fee E V
 79

SOLANACEAE VERBENACEAE
Lycianthes purpusii (Brandegee) Bitter V 1 Aegiphila costaricensis Mold. T 36 (2)
Solandra nitida Zucc. E R Citharexylum affine D. Don T 2

STAPHYLEACEAE VIOLACEAE
Turpinia occidentalis (Swartz) G. Don T 19 (5) Orthion oblanceolatum Lundell T 85 (7)
Rinorea guatemalensis (S. Watson) Bartlett T 68 (1)
TILIACEAE
Heliocarpus donnell-smithii Rose T 1 VITACEAE
Mortoniodendron guatemalense Standl. & Cissus gossypiifolia Standl. V 6
Steyerm. T 35 (6)
Trichospermum mexicanum (DC.) Baill. T 2 (2) VITTARIACEAE
Vittaria lineata (L.) J.E. Smith E R
ULMACEAE
Ampelocera hottlei (Standl.) Standl. T 13 (2) VOCHYSIACEAE
Celtis iguanaeus (Jacq.) Sarg. T 1 Vochysia guatemalensis Donn. Sm. 1 (1)

URTICACEAE ZINGIBERACEAE
Myriocarpa Iongipes Liebm. S 49 (1) Costus spicatus Sw. H 21
Urera caracasana (Jacq.) Griseb. T 1 Costus villosisimus Jacq. H 20
Urera elata (Sw.) Griseb. S 1 Renealmia mexicana Klotzch ex. O.G. Peter-
sen H 11

A P P E N D I X II
Studies on Tropical Lowland Rain Forests, used for comparison of structure and diveristy.

location and classifi- physical characteristics survey characteristics c author

number used cation b
in figures a elevation temperature precipitation dry season # of plot size DBH
(m) (°C) (mm) (months) plots (ha) limit
(cm)

AMERICA
1 Mexico HiEF < 100 25.3 - 26.2 2 2 0 0 - 2700 3- 4 4 0.2 3.3 Sarukhan (1968)
2 Mexico HiEF 350 24.6 2609 3 4 0.25 3.3 Meave del Castillo
(1983)
3 Puerto Rico TRF 500 22.3 3760 1 0.72 4.0 Crow (1980)
4 Costa Rica TMoF 380, 350 24.5, 25.0 2400, 2000 3-4 2 1.2, 0.3 10.0 Holdridge et al.
(1971) d
5 Costa Rica TWF 40-420 25.7-27.5 4 3 0 0 - 5700 0-1 5 0 . 2 - 0.7 10.0 Holdridge et al.
(1971) e
6 Costa Rica TWF < 150 - 3991 0-2 3f 4, 2, 4 10.0 Hartshorn (1982)
7 Panama TF <160 21-32 2670 4 13 0.8-2.0 2.5 Knight (1975)
8 Venezuela DeEF 2 5 0 - 500 - 2000 - lg 155.5 10.0 Rollet (1978, 1979)
9 Venezuela TRF 119 26 3500 0 3 1.0, 0.5 10.0 Uhl & Murphy
(1981)
10 Br. Guyana RF < 300 25.9 2670 3+ 3 5 1.49 10.0 Davis & Richards
(1933/34)
11 Fr. Guyana THu-
DeF < 200 26.0 3450 3 1 1.0 1.0 Lescure et al. (1982)
12 Surinam HiTi-
FiF _ h h 2.0 Schulz (1960)
80

13 E c u a d o r TLF <400 - - 4 0.045 10.0 G r u h b et al. (1963)

14 Brazil TLRF < 100 - - 1 2.0 10.0 C a i n et al. (1956)
15 Brazil TiFiF 40, 60 - - - 6 0.06-0.08 1.0 Takeuchi (1961)
16 Brazil HiTi-
FiF < 100 - - 1 1.0 5.0 P r a n c e et al. (1976)
17 Brazil TiFiF - - - 1 3.5 10.0 Pires et al. (1953)
18 Brazil TiFiF - - - 1 1.0 10.0 Black et al. (1950)
19 Brazil TMoF - 24-25 1200-1800 1-3 i i 10.0 M o r i et al. (1983)

AFRICA
20 G h a n a WEF - ( max 31-33 >1750 - 12 0.0625 10.0 Hall & Swaine
(1981)
21 G h a n a MoEF - ~ min 19-21 1500-1750 - 28 0.0625 10.0 H a l l & Swaine
(1981)
22 Nigeria MiRF < 360 25.5 2080 3 3 1.49 10.0 R i c h a r d s (1939)
23 Nigeria TRF < 76 26.5 1 9 9 0 - 2740 5 4 46.5 10.0 Jones (1955)
24 C a m e r o o n MiRF - 25.5 3 0 0 0 - 3500 3 1 1.49 10.0 R i c h a r d s (1963)

ASIA
25 M a l a y s i a THiF - - - 1 16.0 10.0 Wan Razali (1980)
26 M a l a y s i a LDiF 45-75 25.8 2017 2+3 12 2.04J 30.0 Poore(1968)
27 M a l a y s i a MiLRF 100-200 - - - Ik 1.49 20.0 R i c h a r d s (1936)
28 M a l a y s i a PLRF < 300 m a x 3 2 - 34 5 0 9 0 - 5700 3 4 1.0 10.0 P r o c t o r et aL 0 9 8 3 )
min 2 0 - 2 3
29 M a l a y s i a TRF < 175 - - 1 62.7 40.0 C o u s e n s (1958)
30 T h a i l a n d ERF - - 2696 3 2 0.16 4.5 O g a w a et al. (1965)
31 I n d o n e s i a LDiF 50 - 2347 0 1 1.6 10.0 K a r t a w i n a t a et al.
(1981)
32 I n d o n e s i a TRF . . . . 1 1.0 10.0 D i l m y (1958)
33 New G u i n e a TRF 600-1125 - 2380 - 4 0.8 10.0 P a i j m a n s (1970)

a Figures 7, 8 a n d 9
b De = Dense, Di = Dipterocarp, E = Evergreen, F = Forest, Fi = Firme, H i = H i g h , H u = H u m i d , L = Lowland,
Mi = Mixed, M o = Moist, P = Primary, T = Tropical, Ti = Tierra (Terra), R = Rain, W = Wet
c o n l y plots a n d limits used for c o m p a r i s o n
d plots 20a a n d 23 or t r o p i c a l m o i s t forest
e plots 8a, 8f, 19a, 19b, 19f o f h'opical wet forest
f plots 1, l i b a n d 111, la Selva
g a 25 m wide transect
h at various sites w i t h various plots o f various sizes
i plotless m e t h o d used, total area covered 112 h a
J in a grid, b u t spaced
k o n l y mixed forest plot