516

BIOTROPICA 34(4): 516533 2002

Distribution and Diversity of Pteridophytes and Melastomataceae
along Edaphic Gradients in Yasun National Park,
Ecuadorian Amazonia
1
Hanna Tuomisto
2
, Kalle Ruokolainen
Department of Biology, University of Turku, FIN-20014 Turku, Finland
Axel Dalberg Poulsen, Robbin C. Moran
3
Department of Systematic Botany, Aarhus University, Nordlandsvej 68, DK-8240 Risskov, Denmark
Catalina Quintana, Gustavo Can as, and Jorge Celi
Departamento de Ciencias Biolo gicas, Herbario QCA, Ponticia Universidad Cato lica del Ecuador, Av. 12 de
Octubre # 1076 y Carrio n, Apartado 17012184, Quito, Ecuador
ABSTRACT
We documented the oristic composition of pteridophytes (ferns and fern allies) and Melastomataceae in Yasun
National Park, Amazonian Ecuador. Our main questions were: (1) Are the density of individuals, species richness,
and/or species diversity (measured with Shannons H) of the two plant groups related to edaphic differences? and (2)
How many of the pteridophyte and Melastomataceae species are non-randomly distributed in relation to a soil base
content gradient within terra rme (non-inundated forest). To answer these questions, we sampled 27 line transects
of 500 5 m distributed in an area of ca 20 25 km. The study area included a permanent 50 ha plot established
to monitor forest dynamics; thus, our results also provide information on landscape-scale oristic variability to which
results from within the plot can be compared. A total of 45,608 individuals and 140 species of pteridophytes, and
4893 individuals and 89 species of the Melastomataceae, were counted in the transects. Both with pteridophytes and
with Melastomataceae, a clear negative correlation was found between the amount of extractable bases in the soil and
the number of plant individuals encountered in a transect. With Melastomataceae, species richness and species diversity
also were negatively correlated with soil base content, but with pteridophytes they were not. More than 50 percent
of the common species of both pteridophytes and Melastomataceae were nonrandomly distributed along the soil cation
content gradient within terra rme. We conclude that while the species richness patterns observed in one plant group
are not indicative of similar patterns in other plant groups, it seems likely that a substantial (but unknown) proportion
of species belonging to other plant groups will be found to show distribution patterns that reect edaphic preferences
within terra rme forests.
RESUMEN
Se estudio la composicion or stica de pteridotas (helechos y plantas anes) y Melastomataceas en el Parque Nacional
Yasun , Amazon a ecuatoriana. Nuestras preguntas principales fueron: (1) estan relacionadas la densidad de individuos,
la riqueza de especies y/o la diversidad (medida con la H de Shannon) de los dos grupos de plantas con las diferencias
edacas? y (2) cuantas de las especies de pteridotas y Melastomataceas estan distribuidas en tierra rme de una
manera que no sea al azar, sino en relacion a un gradiente de concentracion de cationes en el suelo? Para desarrollar
estas preguntas, muestreamos 27 transectos de 500 por 5 m distribuidos en una area de 20 por 25 km. El area de
estudio incluye una parcela permanente de 50 ha, establecida para estudiar la dinamica del bosque; por lo tanto,
nuestros resultados proveen informacion sobre la variabilidad or stica en una escala del paisaje que puede compararse
con la de la parcela. En total se encontraron en los transectos 45,608 individuos y 140 especies de pteridotas, y
4893 individuos y 89 especies de Melastomataceas. Tanto las pteridotas como las Melastomataceas, mostraron una
correlacion negativa signicativa entre la cantidad de cationes en el suelo y el numero de individuos de plantas dentro
de un transecto. En las Melastomataceas, la riqueza de especies y la diversidad estuvieron negativamente correlacionadas
con el contenido de cationes en el suelo, pero en las pteridotas, no existio tal correlacion. Mas de 50 por ciento de
las especies comunes de pteridotas y Melastomataceas estuvieron distribuidas dentro de la tierra rme de una manera
no aleatoria en relacion con el gradiente de cationes en el suelo. Concluimos que aunque los patrones de diversidad
de las especies observadas en un grupo de plantas no son indicativos de patrones similares en otros grupos, es probable
que una proporcion apreciable (pero desconocida) de especies pertenecientes a otros grupos de plantas pueda mostrar
patrones de distribucion que reejen preferencias edacas dentro de los bosques de tierra rme.
Key words: Amazonia; diversity; Ecuador; edaphic gradient; Melastomataceae; pteridophytes; species richness; tropical rain
forest.
1
Received 16 December 2001; revision accepted 7 August 2002.
2
e-mail: hantuo@utu.
2
Present address: New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.
Pteridophytes and Melastomataceae in Yasun 517
LOCAL PLANT SPECIES RICHNESS (ALPHA DIVERSITY) has
been suggested to vary along edaphic gradients
within the tropical rain forest realm. Extreme hy-
drological conditions, such as seasonal inundation
or permanent waterlogging, have usually been as-
sociated with fewer tree species per unit area than
well-drained conditions in non-inundated or terra
rme areas (Campbell et al. 1986; Balslev et al.
1987; Duivenvoorden 1994, 1996; Clinebell et al.
1995; Duque et al. 2001; Grandez et al. 2001; Ro-
mero-Saltos et al. 2001).
Within the terra rme areas, however, the pat-
terns of species richness variation are still unknown.
Several studies have addressed species richness
trends along a gradient of soil nutrient content, but
the results have been controversial. Some studies
have indicated that species richness increases with
soil fertility (Gentry 1988; Duivenvoorden 1994,
1996; Tuomisto & Poulsen 1996; Tuomisto 1998),
while others have found exactly the opposite (Hus-
ton 1980), a peak at intermediate fertility (Ashton
1992), or no relationship at all (at least if the ex-
tremely poor soil habitats are excluded; Clinebell et
al. 1995).
Another question with no denite answer is
how different plant species are distributed along
edaphic gradients within tropical forests. Some
well-known cases exist of species with a clear dis-
tribution pattern in relation to extreme conditions,
such as permanent waterlogging or extremely nu-
trient-poor soil (e.g., Mauritia exuosa L.f., which
forms extensive swamp forests in Amazonia, and
the Passiora vitifolia complex discussed in Gentry
1981).
Plant species distributions in the temperate and
boreal zones are known to be strongly affected by
soil characteristics. Hence to understand species
distribution patterns in tropical forests, it is logical
to study whether or not similar relationships exist
there as well. Few studies, however, have explicitly
documented the distribution of Neotropical plant
species in relation to nutrient contents in the soil.
Duivenvoorden (1995) used canonical correspon-
dence analysis (CCA) to study the distribution of
tree species in relation to drainage, ooding, and
soil nutrient content in an area that covered a wide
range of variability in all of these factors. A simple
classication of pteridophyte species according to
their occurrence in study sites with different soil
cation contents was presented by Tuomisto and
Poulsen (1996). This study was restricted to non-
inundated areas, but it included white sand soils
and hence covered a rather long nutrient gradient.
A similarly long gradient was covered in the study
by Tuomisto et al. (1998), which was restricted to
a single fern genus. Often studies dealing with the
responses of plant species to edaphic factors have
used coarse soil classications (Clark et al. 1998,
1999) rather than directly measured chemical
properties. The question remains, therefore, how
plant species are distributed in areas in which dif-
ferences in soil nutrient contents are less pro-
nounced than in the studies cited above.
The present study was conducted in Yasun Na-
tional Park, Amazonian Ecuador. The study area
included both non-inundated and inundated areas,
of which the non-inundated areas (terra rme) have
traditionally been considered homogeneous (i.e.,
without any edaphically differentiated forest types).
In a parallel paper (Tuomisto et al. in press), we
show that changes in species composition of pte-
ridophytes and Melastomatacae (the latter are pre-
dominantly understory trees, shrubs, and vines) in
the study area are associated with differences in
such factors as the content of exchangeable bases
in the soil, presence or absence of inundation, and
topographic heterogeneity within a transect.
In the present paper, our main questions are
the following: (1) Are the density of individuals,
species richness, and/or species diversity of the two
plant groups related to edaphic differences? and (2)
How many of the pteridophyte and Melastomata-
ceae species are nonrandomly distributed in rela-
tion to the soil base content gradient. The edaphic
gradients considered in the present study are rela-
tively short compared to those in earlier studies.
Thus, instead of representing a special case of ex-
treme landscape-scale variability, our results can be
considered relevant for large areas in western Ama-
zonia, where gradients of this magnitude can be
expected to be common at the landscape to re-
gional scales.
Our results also have importance in another
context. Yasun National Park has recently become
an important study site for tropical botany, with
such long-term research activities as the Forest Dy-
namics Project that is currently inventorying a per-
manent 50 ha tree plot in the area. The rst ac-
counts of the oristic composition in subsamples
of the 50 ha plot have already been published (Ro-
moleroux et al. 1997, Svenning 1999, Valencia et
al. in press), and a recent study has inventoried the
woody ora of ve 0.1 ha plots in the surroundings
of the 50 ha plot (Romero-Saltos et al. 2001). Our
study provides additional oristic information from
the area.
518 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
FIGURE 1. Location map of the study area in Ama-
zonian Ecuador. The transects are identied with the
numbers used in the text. The rectangle next to transect
21 is a permanent 50 ha forest dynamics plot.
In the present study, we documented the o-
ristic composition of pteridophytes and Melasto-
mataceae in the park, using 27 line transects dis-
tributed in an area of ca 20 25 km that included
the 50 ha plot. Our objective was to characterize
the overall species composition of the target groups
in the area, which can later be used as a regional
data set against which the results from within the
50 ha plot can be compared.
METHODS
FIELDWORK. Fieldwork was carried out in the vi-
cinity of the Yasun Research Station, situated in
Yasun National Park and operated by Ponticia
Universidad Catolica del Ecuador. The sampled
sites were between 032S and 046S and from
7621W to 7632W, in an area of ca 25 20
km (Fig. 1). The area is accessible by a road con-
structed for oil drilling, and access is presently con-
trolled by the oil company to prevent colonization
and agricultural expansion within the national
park.
Yasun National Park lies 200300 m above sea
level, and the climate is tropical, humid, and al-
most aseasonal. Annual precipitation exceeds 2400
mm, and no month receives less than 100 mm of
rain on average, although drier months may occur
in some years (Canadas 1983, Balslev et al. 1987).
Vegetation in the study area consists mainly of
closed-canopy, non-inundated terra rme forests.
The terra rme areas are interrupted by river ood-
plains with seasonally inundated forests and by
swamps dominated by the palm Mauritia exuosa.
Soil and oristic studies were conducted in 27
sites using transects 500 5 m (Fig. 1). The ma-
jority of the transects were placed in non-inundated
forests, but three were placed in seasonally inun-
dated areas for comparison. Transects 6 and 22
were situated in the oodplain of the Tiputini Riv-
er, and transect 13 was situated in the oodplain
of a creek. In the middle of each transect, a narrow
trail (baseline) was opened that followed a prede-
ned compass bearing. For practical reasons, most
transects were started near the road, from which
they were directed deeper into the forest to keep
them in undisturbed vegetation.
The topography of each transect was measured
along the baseline using a clinometer (Suunto, Fin-
land). The difference in elevation between the
highest and lowest point in a transect was used as
a simple measure of topographic heterogeneity,
which was expected to correlate with variation in
moisture availability and other soil differences that
are related to catena formation.
Surface soil samples (the top 5 cm of the min-
eral soil) were collected near the beginning, middle,
and end of each transect. Each of the soil samples
consisted of ve pooled subsamples collected with-
in an area of ca 5 5 m. Details of the soil analyses
(both methodology and results) will be published
elsewhere (Tuomisto et al. in press). In the present
paper, only information on the total concentration
of extractable bases (Ca, K, Mg, and Na analyzed
in 1 M NH
4
OAc at pH 7; van Reeuwijk 1993)
will be used.
Pteridophytes (ferns and fern allies) and mem-
bers of the family Melastomataceae (excluding Me-
mecylaceae) were identied to species and counted
within an estimated 2.5 m at both sides of the
baseline for each transect, yielding a sample size of
0.25 ha per transect. To expedite eldwork, the
lateral limits of the transect were not marked, but
the distance between the baseline and borderline
plant individuals were occasionally measured to
maintain consistency in transect width.
For pteridophytes, only individuals with at least
one leaf longer than 10 cm were recorded, and
epiphytic and climbing individuals were included
only if they had green leaves at a height less than
2 m above ground. Because earlier studies differ in
whether epiphytic pteridophytes were included
(Tuomisto & Ruokolainen 1994, Tuomisto et al.
1995, Ruokolainen et al. 1997) or excluded (Poul-
sen & Balslev 1991; Tuomisto & Poulsen 1996,
2000), we included them but counted ground pte-
ridophytes (self-supporting individuals with rhi-
zomes in the soil) separately from individuals that
were either epiphytic or climbing on a tree trunk
Pteridophytes and Melastomataceae in Yasun 519
or log. Results will be presented for the data set
containing all individuals, except when explicitly
stated otherwise. For vegetatively spreading species
such as Selaginella, each rooting stem was counted
as one individual.
All individuals of Melastomataceae, including
seedlings that had at least one post-cotyledon leaf,
were recorded. Voucher collections for all species
of both plant groups are deposited in herbaria in
Ecuador (QCA, QCNE), Denmark (AAU), Fin-
land (TUR), and the United States (NY). Unicates
are kept in QCA.
DATA ANALYSES. Chi-square tests were conducted
to determine how many of the pteridophyte and
Melastomataceae species were distributed in a non-
random way across the observed gradient of soil
cation content. Only the 24 terra rme transects
were used in these analyses to avoid the confound-
ing effect of inundation. For the rst set of chi-
square analyses, the transects were allocated to two
groups: low and high soil base content (N 12 in
both groups; base contents 1.166.15 cmol()/kg
and 8.9727.3 cmol()/kg, respectively). For the
second set of analyses, transects were allocated to
three groups: low, intermediate, and high soil base
contents (N 8 in each group; base contents
1.164.81 cmol()/kg, 4.8912.5 cmol()/kg,
and 12.727.3 cmol()/kg, respectively). The ra-
tionale for grouping the transects in two different
ways was that the two-group test allowed identi-
fying species for which occurrences were concen-
trated to either half of the cation gradient, while
the three-group test made it possible to identify
species with occurrences that were concentrated to-
ward the middle of the gradient or to its either
extreme. Only species that occurred in at least 4 of
the 24 transects were tested.
A standard chi-square test assumes that each
plant individual is statistically independent of other
individuals and yields one degree of freedom when
testing whether the corresponding species shows a
nonrandom distribution with respect to the group-
ing criterion. We reasoned that this assumption was
not warranted because individuals within the same
transect were probably highly autocorrelated.
Therefore, we estimated the statistical signicance
of each test with a Monte Carlo permutation pro-
cedure using transects as the permutable units in-
stead of individuals: the probability of error of the
observed chi-square statistic was estimated by com-
paring its value to 999 values obtained by permut-
ing transect positions along the cation gradient
while keeping the transect-wise plant abundances
xed. Spatial autocorrelation among transects was
not considered a problem because Mantel tests be-
tween the patterns of geographical and oristic dis-
tances among the transects did not show signicant
correlations (Tuomisto et al. in press). The program
Resampling Stats 4.1 was used for the chi-square
tests.
Pearson correlation coefcients were calculated
to test for relationships of individual density, spe-
cies richness, and diversity with soil cation content
and topographic heterogeneity. The correlation co-
efcients were calculated with the program
CorrPermute (available at http://www.fas.umontreal.
ca/BIOL/Legendre/indexEnglish.html), which allows
estimation of the statistical signicance of r with a
Monte Carlo permutation test; 999 permutations were
used.
RESULTS
PTERIDOPHYTES. A total of 45,608 individuals and
140 species of pteridophytes were counted in the
27 transects. Of these, 33,769 individuals (74% of
all individuals) and 108 species (77% of all species)
were found growing on the ground. Because three
species pairs were confused in the eld, they have
been lumped for the numerical analyses, which
therefore only deal with 106 species of ground pte-
ridophytes and a total of 137 species (Table 1).
The total number of pteridophyte individuals
found within one transect varied between 734 and
3338, the mean being 1689 individuals per transect
(6760 individuals/ha). The number of pterido-
phyte individuals on the ground varied between
360 and 2521 per transect, the mean being 1251
(5004 individuals/ha). The number of species per
transect varied between 21 and 65 for all pterido-
phytes (x 47), and between 12 and 46 for
ground pteridophytes (x 35).
The most abundant pteridophyte was Adian-
tum humile Kunze, which was represented by 5000
individuals in 26 transects (Table 1). The other
species with more than 2000 individuals were Poly-
botrya osmundacea Willd., P. crassirhizoma Lellinger,
and Lomariopsis japurensis (Martius) J. Smith. Only
two species were found in all 27 transects, namely
L. japurensis and A. obliquum Willd. Twenty-six
species were found in only one transect, and 11 of
these were represented by a single individual. The
mean number of individuals per species was 330,
and the mean frequency (number of transects in
which the species was found) was 9.
Out of the ten most abundant pteridophyte
species, ve were signicantly most abundant in
520 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
TABLE 1. Pteridophyte species found in 27 transects in Amazonian Ecuador. A representative voucher specimen is indicated
for each species (collection numbers from the series of R. C. Moran). The general life-form is given for each
species: t terrestrial; sc scandent; cl climber; l lithophyte; and e epiphyte. The total number of
individuals found in each species is shown, as well as the number of terrestrial and scandent individuals as
opposed to epiphytic, epilithic, and climbing individuals. Frequency is the number of transects in which a species
was encountered. Three species pairs were inconsistently identied in the eld and are therefore lumped in the
table and in the numerical analyses ( * Antrophyum cajenense and A. guayanense Hieron.; ** Bolbitis
serratifolia and Bolbitis sp. 2; *** Danaea acuminata and D. oblanceolata Stolze).
Species
Number of individuals
Coll.
number Life-form
On
ground
On other
substrates Total
Fre-
quency
Adiantum cayennense Klotzsch
A. humile Kunze
A. latifolium Lam.
A. macrocladum Klotzsch
A. cf. olbiquum Willd.
6238
6237
6116
6256
6043
t
t
t
t
t
10
4978
11
12
1596
22
12
10
5000
11
12
1608
5
26
3
3
27
A. petiolatum Desv.
A. platyphyllum Sw.
A. pulverulentum L.
A. scalare R. M. Tryon
A. terminatum Kunze ex Miq.
6186
6212
6159
6255
6050
t
t
t
t
t
22
3
1974
1
96
1
1
22
3
1975
1
97
7
1
26
1
8
A. wilsonii Hook.
Adiantum sp. 1
Adiantum sp. 2
Alsophila cuspidata (Kunze) D. S. Conant
6181
6138
6239
6049
t
t
t, (e)
t
22
19
36
209
7
22
19
43
209
1
3
3
25
Anetium citrifolium (L.) Splitgb.
Antrophyum cajenense (Desv.) Sprengel*
Asplenium angustum Sw.
A. auritum Sw.
A. cirrhatum Willd.
A. cuneatum Lam.
A. juglandifolium Lam.
6082
6052
6074
6041
6136
6268
6103
e
e
e
e
e, (t)
e
e, t
76
4
72
37
3
73
377
5
3
72
37
3
73
453
5
7
23
15
1
11
22
2
4
A. ortegae Murakami and R. C. Moran
A. pearcei Baker
A. repandulum Kunze
A. serratum L.
A. stuebelianum Hieron.
Asplenium sp. 1
Bolbitis lindigii (Mett.) C. Chr.
B. nicotianifolia (Sw.) Alston
B. oligarchica (Baker) Hennipman
B. serratifolia (Kaulf.) Schott**
6146
6134
6228
6269
6077
6229
6046
6112
6244
6141
t, (e)
e, (t)
t
e
e, (t)
t
cl
cl
t
t, l
862
5
1
59
87
564
512
542
25
38
176
17
468
244
291
5
11
900
181
1
17
527
87
808
803
547
36
18
22
1
10
22
2
20
7
15
2
Campyloneurum cf. abruptum (Lindm.)
B. Leon
Campyloneurum aphanophlebium (Kunze)
T. Moore
C. cf. brevifolium (Link) Link
C. chlorolepis Alston
6177
6185
6113
6114
e
e
e
e
15
14
1
3
15
14
1
3
3
3
1
3
C. fuscosquamatum Lellinger
C. phyllitidis (L.) C. Presl
Campyloneurum sp. 1
Ctenitis refulgens (Mett.) Vareschi
Cyathea amazonica R. C. Moran
C. lasiosora (Mett. ex Kuhn) Domin
C. pungens (Willd.) Domin
6065
6242
6233
6192
6045
6079
6111
e
e
t, e
t
t
t, (e)
t, (e)
4
14
2
38
237
380
726
11
7
17
21
730
11
21
2
38
254
401
25
10
1
2
6
14
20
C. trichiata (Maxon) Domin
Cyclodium trianae (Mett.) A. R. Sm.
Cyclopeltis semicordata (Sw.) J. Sm.
6162
6047
6132
t
t
t
3
2
132
3
2
132
2
1
5
Danaea acuminata H. Tuomisto and
R. C. Moran***
D. bipinnata H. Tuomisto
6209
6260
t
t
274
37
274
37
19
1
D. falcata H. Tuomisto and
R. C. Moran
6259
6064
t
t
23
297 1
23
298
1
22
Pteridophytes and Melastomataceae in Yasun 521
TABLE 1. Continued.
Species
Number of individuals
Coll.
number Lifeform
On
ground
On other
substrates Total
Fre-
quency
D. nodosa (L.) Sm.
D. cf. ulei H. Christ
Dennstaedtia bipinnata (Cav.) Maxon
Didymochlaena truncatula (Sw.) J. Smith
Diplazium bombonasae Rosenst.
6046
6184
6130
6032
6253
t
t
t, (e)
t, (e)
t, (e)
297
70
7
1093
297
1
1
99
7
298
70
8
1192
304
22
7
4
25
4
D. grandifolium (Sw.) Sw. var.
andicola Stolze
D. pinnatidum Kunze
D. tungurahuae (Sodiro) C. Chr.
6235
6029
6158
t
t
t
83
1461
734 1
83
1461
735
11
22
14
Elaphoglossum accidum (Fee) T. Moore
E. luridum (Fee) H. Christ
E. cf. megalurum Mickel
E. nigrescens (Hooker) Diels
6227
6179
6262
6243
e
e
e
e
161
31
1
45
161
31
1
45
11
8
1
7
E. raywaense (Jenm.) Alston
Elaphoglossum sp. 1
Hemidictiym marginatum (L.) C. Presl
6153
6206
6163
e
e
t, (e) 6
90
3
2
90
3
8
16
1
4
Lindsaea cf. divaricata Klotzsch
L. lancea (L.) Bedd. var. lancea
L. phassa K. U. Kramer
Lindsaea sp. 1
Lomagramma guianensis (Aubl.) Ching
Lomariopsis fendleri D. C. Eaton
L. japurensis (Mart.) J. Smith
L. latipinna Stolze
L. nigropaleata Holttum
Lygodium volubile Sw.
6257
6280
6183
6211
6048
6129
6133
6027
6053
6097
t
t, (e)
t
t
cl
e, (cl)
cl
cl
cl
sc
9
4
1
31
464
129
7
19
7
610
26
1880
505
789
9
4
1
31
1074
26
2009
512
808
7
3
1
1
6
16
9
27
19
13
3
Microgramma fuscopunctata (Hook.)
Vareschi
M. percussa (Cav.) Sota
Nephrolepis pendula (Raddi) J. Sm.
6071
6220
6083
e
e
e, (t) 4
361
2
15
361
2
19
25
2
9
N. rivularis (Vahl.) Mett. ex Krug 6279 e, (t) 30 30 4
Pecluma hygrometrica (Splitg.)
M. G. Price 6247 e 20 20 8
P. ptilodon (Kunze) M. G. Price var.
pilosum (A. M. Evans) Stolze
Pityrogramma calomelanos (L.) Link
6246
6273
e
t 1
7 7
1
7
1
Pleopeltis macrocarpa (Willd.) Kaulf. var.
laciniata Stolze
Polybotrya caudata Kunze
P. crassirhizoma Lellinger
P. fractiserialis (Baker) J. Sm.
6161
6060
6066
6125
e
cl
cl
cl
1103
778
37
1
554
1574
5
1
1657
2352
42
1
26
26
4
P. osmundacea Willd.
P. pubens Mart.
Polypodium adnatum Klotzsch
P. bombycinum Maxon
P. caceresii Sodiro
P. dasypleuron Kunze
P. decumanum Willd.
Polypodium sp. 1
Pteris altissima Poiret
P. pungens Willd.
Saccoloma inaequale (Kunze) Mett.
Salpichlaena volubilis (Kaulf.) J. Sm.
6030
6063
6258
6037
6051
6092
6080
6119
6275
6266
6054
6034
cl
cl
e
e
e
e
e
e
t
t
t
sc
1045
105
1
2
4
317
126
1785
84
11
8
38
38
8
1
3
1
2830
189
11
8
39
38
8
1
2
4
320
127
25
5
1
2
14
11
4
1
2
2
21
17
Selaginella anceps (C. Presl) C. Presl
S. exaltata (Kunze) Spring
S. geniculata (C. Presl) Spring
S. haematodes (Kunze) Spring
S. lechleri Hieron.
6170
6109
6207
6223
6276
t, (e)
sc
t
t, e
t
1109
382
968
65
11
120
4
59
1229
382
972
124
11
7
17
17
7
1
522 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
TABLE 1. Continued.
Species
Number of individuals
Coll.
number Lifeform
On
ground
On other
substrates Total
Fre-
quency
S. parkeri (Hook. and Grev.) Spring
S. revoluta Baker
S. speciosa A. Br.
6078
6101
6226
t
t
t, (e)
908
9
125
2
53
910
9
178
5
3
5
Stigmatopteris heterophlebia (Baker)
R. C. Moran
S. opaca (Baker) C. Chr.
Tectaria cf. antioquoiana (Baker) C. Chr.
6166
6126
6105
t
t
t
227
38
1911 3
227
38
1914
13
5
18
T. draconoptera (D. C. Eaton) Copel.
T. pilosa (Fee) R. C. Moran
T. plantaginea (Jacq.) Maxon
6069
6189
6173
t
t
t
96
56
33
96
56
33
12
7
3
T. pubens R. C. Moran 6180 t 169 169 7
T. cf. vivipara Jermy and
T. G. Walker
Tectaria sp. 1
Tectaria sp. 2
Thelypteris abrupta (Desv.) Proctor
6190
6148
6104
6187
t
t
t
t
505
228
1252
33
2
12
507
228
1264
33
8
18
23
9
T. ancyriothrix (Rosenst.)
A. R. Sm. 6215 t 13 13 5
T. arcana (Maxon and C. V. Morton)
C. V. Morton 6198 t 1 1 1
T. biformata (Rosenst.) R. M. Tryon 6107 t, (e) 490 11 501 14
T. dentata (Forssk.) E. St. John 6140 t, e 3 2 5 5
T. glandulosa (Desv.) Proctor
var. brachyodus (Kunze) A. R. Sm.
T. hispidula (Decne.) C. F. Reed
T. juruensis (C. Chr.) R. M. Tryon and
D. S. Conant
T. leprieurii (Hook.) R. M. Tryon
var. incana (H. Christ) A. R. Sm.
6218
6310
6241
6312
t
t
t
t
199
2
838
5
3
199
2
841
5
3
1
6
2
T. macrophylla (Kunze) C. V. Morton
T. opposita (Vahl) Ching
T. opulenta (Kaulf.) Fosberg
T. pennata (Poir.) C. V. Morton
T. tristis (Kunze) R. M. Tryon
Thelypteris sp. 1
Thelypteris sp. 2
Trichomanes collariatum Bosch
6040
6311
6139
6028
6143
6201
6142
6031
t, (e)
t
e, t
t
t
t
t
cl
1752
1
24
483
11
205
2
28
39
3
21
1780
1
63
486
11
205
2
21
15
1
12
17
3
3
1
6
T. diversifrons (Bory) Mett. 6067 t 245 245 21
T. hostmannianum (Klotzsch) Kunze
T. pinnatum Hedwig
T. polypodiodes L.
Trichomanes sp. 1
Triplophyllum funestum (Kunze) Holttum
6087
6264
6287
6265
6090
t
t
e
t
t
58
133
10
55
1
58
133
1
10
55
1
15
1
1
7
137 spp. recognized in numerical analyses
(total 140 spp.)
33,769 11,839 45,608
the transects in which soils had a lower than me-
dian exchangeable base content. Two species
showed a statistically signicant bias away from the
poorest soils, while the distribution of three species
was statistically indistinguishable from random
along the soil gradient (Fig. 2). Two of the abun-
dant species, Polybotrya caudata Kunze and L. ja-
purensis, were especially abundant in transects 6
and 22, which were affected by the inundations of
the Tiputini River. Most of the other species were
either rare or absent in these transects. Of course,
even the terra rme transects often included poorly
drained sections in places where the transect
crossed a creek or a poorly drained valley, but as
the present analyses were done at the level of entire
transects, distributions at ner spatial scales were
not assessed.
Out of the 84 pteridophyte species that oc-
Pteridophytes and Melastomataceae in Yasun 523
FIGURE 2. The abundances (numbers of individuals) of the ten most abundant pteridophyte species (left column)
and the ten most abundant Melastomataceae species (right column) in 27 transects in Amazonian Ecuador. The
transects are arranged in increasing order of soil cation content (sum of exchangeable Ca, K, Mg, and Na). Total
number of individuals of each species is given in parentheses after the species name. Bars corresponding to terra rme
transects are black; those corresponding to transects affected by inundations are gray.
curred in at least 4 of the 24 terra rme transects,
43 (51%) were signicantly (P 0.05) biased in
their distribution among the transect groups that
had been dened according to soil base content
(i.e., they yielded a signicant result in the two-
group test or the three-group test, or both). Chi-
square tests gain power when frequencies increase,
but there was no obvious trend in the percentage
of species with biased distributions when the least
frequent species were successively excluded; the
percentage varied but remained between 50 and 60
percent (Fig. 3A).
A clear negative correlation was found between
the amount of extractable bases in the soil and the
524 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
FIGURE 3. Percentage of species of pteridophytes (A) and Melastomataceae (B) that show statistically signicant
(chi-square test, P 0.05) biases in their distribution among two or three groups of non-inundated transects as a
function of the minimum number of transects in which the species were observed. The 24 transects were allocated
to the groups on the basis of the concentration of extractable bases in the soil. The number of species that occurred
in at least four transects is 84 for pteridophytes and 59 for Melastomataceae. The percentages corresponding to the
higher minimum numbers of transects are based on fewer species; the curves are truncated at the point where only
10 species are included. The proportion of species expected to show a statistically signicant response by chance is 5
percent both for the two-group test and the three-group test, which is clearly lower than the percentages found here.
number of pteridophyte individuals found in a
transect (r 0.44, P 0.03; Fig. 4A). The num-
ber of species and species diversity (as measured
with Shannons H), however, were uncorrelated
with soil base content (Fig. 4C, E). The same re-
sults were obtained whether all 27 transects were
considered, or only the 24 that were situated in
terra rme. When only ground pteridophytes were
included in similar analyses, none of the correla-
tions were statistically signicant.
The degree of topographic heterogeneity within
the transect was not signicantly correlated with
the number of pteridophyte individuals. The same
result was obtained whether the three seasonally
inundated transects were included in the analysis
or not (Fig. 5A). Both species richness and diversity
were positively correlated with topographic hetero-
geneity, and especially its logarithm, when all tran-
sects were included. This was entirely due to the
three seasonally inundated transects which were
both species-poor and topographically at; no cor-
relation was found when only terra rme transects
were analyzed (Fig. 5C, E). The results obtained
with ground pteridophytes were similar.
MELASTOMATACEAE. A total of 4893 individuals
and 89 species of the Melastomataceae were count-
ed in the 27 transects (Table 2). The number of
individuals found within one transect varied be-
tween 59 and 372, the mean being 181 individuals
per transect (724 individuals/ha). The number of
species per transect varied between 9 and 44, the
mean being 28.
The most abundant Melastomataceae species
was Miconia triplinervis R. & P., which was repre-
sented by 345 individuals in 18 transects (Table 2).
Miconia serrulata (DC.) Naud. was also represented
by more than 300 individuals. Miconia elata (Sw.)
DC. had the highest frequency as it was found in
23 transects. Eleven species were found in only 1
transect and 3 of these were represented by only 1
individual. The mean number of Melastomataceae
individuals per species was 55, and the mean fre-
quency (number of transects in which the species
was found) was 9.
Out of the ten most abundant Melastomata-
ceae species, ve were signicantly most abundant
in the transects in which soils had a lower than
median exchangeable base content. Two species
were signicantly most abundant in the transects
having soils with a higher base content, while the
distribution of three species was statistically indis-
tinguishable from random along the soil gradient
(Fig. 2). Only one of the abundant species, M.
serrulata, was especially abundant in transects 6 and
22, which were affected by the inundations of the
Tiputini River. The other species were either rare
or absent in these transects.
Out of the 59 Melastomataceae species that oc-
curred in at least four of the terra rme transects,
24 (41%) were signicantly biased in their distri-
bution among transect groups dened using soil
Pteridophytes and Melastomataceae in Yasun 525
FIGURE 4. The number of individuals (A, B), number of species (C, D), and species diversity (as measured with
Shannons H; E, F) plotted against the mean concentration of extractable bases in the soil (sum of Ca, K, Mg, and
Na) in each of 27 transects in Amazonian Ecuador. A, C, E: Pteridophytes; B, D, F: Melastomataceae. Round symbols
indicate terra rme transects; squares indicate transects in river or creek oodplains. The regression line for all 27
transects is shown in black; that for the 24 terra rme transects is shown in gray.
base content (i.e., they yielded a signicant result
in the two-group test or the three-group test, or
both). The percentage of species with biased dis-
tributions tended to increase with increasing mi-
numum frequency of the species, until among spe-
cies that had been found in at least ten transects
the percentage more or less stabilized at between
50 and 60 percent (Fig. 3B).
With Melastomataceae, clear negative correla-
tions were found between the amount of extract-
526 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
FIGURE 5. The number of individuals (A, B), number of species (C, D), and species diversity (as measured with
Shannons H; E, F) plotted against the largest elevation difference measured within each of 27 transects in Amazonian
Ecuador. A, C, E: Pteridophytes; B, D, F: Melastomataceae. Round symbols indicate terra rme transects; squares
indicate transects in river or creek oodplains. The best-tting regression curve for all 27 transects is shown in black;
that for the 24 terra rme transects is shown gray. The nonlinear curves are either logarithmic (C, E) or quadratic (B,
D, F).
Pteridophytes and Melastomataceae in Yasun 527
able bases in the soil, and the number of individ-
uals, number of species, and Shannons H diversity
index. The correlation between extractable bases
and individuals was slightly more pronounced
when all 27 transects were considered (r 0.65,
P 0.001) than when only the 24 terra rme
transects were considered (r 0.62, P 0.001;
Fig. 4B). The opposite was true of both the cor-
relations between extractable bases and number of
species (r 0.56, P 0.004 with all transects, r
0.60, P 0.003 with terra rme transects) and
Shannons H (r 0.46, P 0.02 with all tran-
sects, r 0.56, P 0.008 with terra rme tran-
sects; Fig. 4D, F).
The degree of topographic heterogeneity within
the transect was negatively correlated with the
number of Melastomataceae individuals when the
seasonally inundated transects were excluded, but
not when they were included in the analysis. Both
with the terra rme data set and the full data set,
a quadratic regression gave a better t than a
straight line (Fig. 5B). The peak at intermediate
topographic heterogeneity was especially strong
with species richness and diversity, both of which
showed a statistically signicant quadratic response
function but no linear correlation. These results
were similar with the full data set and the 24 terra
rme transects (Fig. 5D, F).
COMPARISON OF THE TWO PLANT GROUPS. The Me-
lastomataceae were clearly less abundant than ter-
restrial pteridophytes; the number of Melastoma-
taceae individuals was only 14 percent of the num-
ber of terrestrial pteridophyte individuals. The dif-
ference in species richness, however, was relatively
small; the number of Melastomataceae species was
82 percent of the number of terrestrial pterido-
phyte species.
There was a signicant positive correlation be-
tween the number of individuals of the two plant
groups in a transect (r 0.64, P 0.003), which
became even more pronounced when the three
transects affected by inundations were excluded (r
0.72, P 0.001; Fig. 6A). With all 27 transects,
there was also a positive correlation with the num-
ber of species found in a transect (r 0.48, P
0.01), but this correlation disappeared when the
three seasonally inundated transects were excluded
(Fig. 6B). The diversity in the transects (as mea-
sured with Shannons H) was not correlated be-
tween the two plant groups no matter whether the
seasonally inundated transects were included or not
(Fig. 6C).
A species optimum for soil base content was
calculated as the weighted mean of the soil base
contents in the transects where the species was pre-
sent, using species abundance as the weight (Jong-
man et al. 1995). Optima were calculated for all
species with at least ten individuals in the data set,
and they ranged from 1.2 to 19.3 cmol()/kg for
pteridophytes and from 3.1 to 19.6 cmol()/kg for
Melastomataceae. Although both the minimum
and maximum values were lower for pteridophytes,
the mean was slightly higher [9.1 cmol()/kg] than
for Melastomataceae [8.5 cmol()/kg]. The fre-
quency distributions of the species optima were
somewhat different between the two plant groups.
Pteridophyte species optima were evenly distribut-
ed along the gradient (although extreme values
were underrepresented), but Melastomataceae spe-
cies optima were strongly skewed toward values be-
low 10 cmol()/kg (Fig. 7).
DISCUSSION
The density of pteridophyte individuals varied
widely in the different transects inventoried in Ya-
sun : 144010,080 individuals/ha for ground pte-
ridophytes and 294013,350 individuals/ha for all
pteridophytes. These values span almost the entire
range of 3390 to 13,930 individuals/ha reported
earlier at eight lowland Amazonian sites in Peru
and Ecuador (Tuomisto & Poulsen 1996). Densi-
ties of the Melastomataceae in Yasun ranged from
240 to 1490 individuals/ha, which seems low when
compared to the 1740 individuals/ha reported at a
site in Peru (Tuomisto & Ruokolainen 1994).
Species richness per transect also varied widely
in Yasun . The mean species richness (35) of
ground pteridophytes in the Yasun transects is
high compared with quantitative survey samples
from elsewhere in lowland Amazonian Ecuador
and Peru (Tuomisto & Poulsen 1996), which in-
cluded 1431 species in 0.25 ha. With the Melas-
tomataceae, the species richness values reported
from Peru, 1422 species/0.35 ha (Tuomisto &
Ruokolainen 1994), fall within the range but below
the mean (28 species/0.25 ha) found in the present
study.
Earlier studies with pteridophytes have shown
that density of individuals, species richness, and di-
versity can vary considerably even at the local scale
(Tuomisto & Poulsen 2000). In some cases, this
variation seemed to be related to topographic var-
iation within a transect, but not always. In the pre-
sent study, the amount of topographic heteroge-
neity within a transect was not correlated with ei-
ther density of individuals, species richness, or di-
528 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
TABLE 2. Melastomataceae species found in 27 transects in Amazonian Ecuador. A representative voucher specimen is
indicated for each species (collection numbers from the series of K. Ruokolainen). The total number of individuals
found in each species is shown. Frequency is the number of transects in which a species was encountered.
Morphospecies numbers are not strictly sequential because the same numbering system is being used in other
study areas, and some of the already numbered morphospecies were not encountered in the present study.
Coll.
number
Number of
individuals Frequency
Aciotis purpurascens (Aubl.) Tr.
Adelobotrys boissieriana Cogn.
Adelobotrys sp. 1
Adelobotrys sp. 2
Adelobotrys sp. 4
Bellucia pentamera Naud.
Blakea bracteata Gleason
B. rosea (R. and P.) D. Don
Clidemia allardii Wurdack
10362
10353
10294
10332
10380
10357
10302
10285
10295
3
7
14
49
12
6
2
60
14
1
1
6
11
6
5
2
21
7
C. dimorphica Macbr.
C. epiphytica (Tr.) Cogn.
C. heterophylla (Desr.) Gleason
C. japurensis DC.
C. septuplinervia Cogn.
C. sprucei Gleason
Clidemia sp. 3
Henriettella sp. 1
10254
10355
10253
10390
10261
10340
10323
10344
139
110
92
7
171
70
22
14
20
22
20
1
22
6
7
2
Leandra aristigera (Naud.) Cogn.
L. caquetana Sprague
L. chaetodon (DC.) Cogn.
L. longicoma Cogn.
L. macdanielii Wurdack
L. secunda (D. Don) Cogn.
L. secundiora (DC.) Cogn.
Leandra sp. 1
Loreya sp. 1
Loreya sp. 2
10378
10319
10381
10266
10301
10280
10315
10325
10333
10389
19
40
6
137
11
15
1
10
2
6
5
16
3
22
2
4
1
6
1
4
Maieta guianensis Aubl.
Miconia abbreviata Markgraf
M. acutipetala Sprague
M. amazonica Tr.
M. ampla Tr.
M. aurea (D. Don) Naud.
M. aureoides Cogn.
M. centrodesma Naud.
M. decurrens Cogn.
M. elata (Sw.) DC.
10289
10268
10347
10352
10379
10349
10368
10371
10366
10258
87
24
17
1
2
18
131
7
49
225
13
12
10
1
2
9
11
3
11
23
M. fosteri Wurdack
M. grandifolia Ule
M. lamprophylla Tr.
M. longifolia (Aubl.) DC.
M. mazanana Macbr.
M. napoana Wurdack
M. nervosa (Smith) Tr.
M. paleacea Cogn.
M. prasina (Sw.) DC.
10270
10278
10339
10305
10312
10343
10290
10296
10331
115
237
14
52
28
48
62
3
13
11
19
3
5
1
19
15
3
6
M. procumbens (Gleanson) Wurdack
M. pterocaulon Tr.
M. punctata (Desr.) D. Don
M. schunkei Wurdack
M. serrulata (DC.) Naud.
M. tomentosa (L. C. Rich.) D. Don
M. trinervia (Sw.) D. Don ex Loud.
M. triplinervis R. and P.
10252
10286
10284
10306
10259
10276
10264
10267
39
6
96
75
307
42
39
345
11
3
17
10
22
7
12
18
Miconia sp. 3
Miconia sp. 4
10256
10320
80
10
21
1
Pteridophytes and Melastomataceae in Yasun 529
TABLE 2. Continued.
Coll.
number
Number of
individuals Frequency
Miconia sp. 7
Miconia sp. 12
Miconia sp. 13
Miconia sp. 14
Miconia sp. 15
Miconia sp. 17
Miconia sp. 18
10374
10334
10382
10311
10372
10304
10265
109
1
3
9
20
116
234
16
1
2
1
5
7
17
Miconia sp. 19
Miconia sp. 20
Miconia sp. 21
Miconia sp. 22
Miconia sp. 23
Miconia sp. 24
Miconia sp. 25
Miconia sp. 26
Miconia sp. 27
10375
10257
10354
10338
10348
10376
10346
10316
10300
80
6
21
6
3
213
60
10
14
16
3
4
6
2
21
13
3
3
Miconia sp. 29
Miconia sp. 32
Miconia sp. 33
Miconia sp. 34
Miconia sp. 35
Miconia sp. 39
Miconia sp. 40
Miconia sp. 41
Miconia sp. 43
10321
10330
10373
10281
10350
10260
10318
10369
10328
54
2
48
8
17
2
144
12
19
18
1
7
2
2
2
11
5
8
Miconia sp. 44
Miconia sp. 47
Monolena primulaeora Hook.
Ossaea boliviensis (Cogn.) Gleason
Tococa caquetana Sprague
T. discolor Pilger
Tococa sp. 1
Triolena amazonica (Pilger) Wurdack
10282
10329
10335
10271
10273
10360
10303
10297
48
25
11
85
103
39
67
93
8
5
2
13
19
5
3
11
Total 89 species 4893
versity within the terra rme area. Especially with
species richness, this result is somewhat surprising,
because increasing topographic heterogeneity
would be expected to provide a wider suite of hab-
itats and hence increase beta diversity within the
transect. The expected pattern was found only
when the three transects of oodplain forests were
included in the analysis, but it is likely that this
result was due to the diversity-decreasing effect of
ooding, rather than topography.
With Melastomataceae, very different patterns
were found in relation to topography than with
pteridophytes. A clear peak at intermediate topo-
graphic heterogeneity was found with density of
individuals, species richness, and species diversity.
Not only the at oodplain transects and some
other relatively at transects showed low species
richness and diversity, but also the transects with
highest topographic heterogeneity. This surprising
result may be due to the tendency of the transects
with most rugged topography to have relatively
base-rich soils, a factor that correlated negatively
with the species richness and diversity of Melasto-
mataceae. Consequently, the topographic variabili-
ty gradient in the present case is a complex gradient
that not only represents a gradient of heterogeneity
in soil moisture, but where the apparent plant re-
sponse may actually be controlled by different lim-
iting factors in the lower end of the gradient than
in the upper end of the gradient (cf. Pausas & Aus-
tin 2001). Of course, our study has only estab-
lished the existence of correlations and not of caus-
al relationships; to unravel the latter, other soil
properties would have to be included as well, pref-
erably in an experimental framework to clarify the
roles of intercorrelated explanatory factors.
It has often been pointed out that local species
richness is not only dependent on local processes,
530 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
FIGURE 6. Correlation between pteridophytes and
Melastomataceae for number of individuals (A), number
of species (B), and species diversity (as measured with
Shannons H; C) in 27 transects in Amazonian Ecuador.
The regression line for all 27 transects is shown in black;
FIGURE 7. Frequency distribution of pteridophyte and
Melastomataceae species optima for different soil base
content intervals in 27 transects in Amazonian Ecuador.
Species represented by less than ten individuals are not
included.

that for the 24 terra rme transects is shown in gray.

Symbols indicate the mean concentration of exchangeable
bases in the soils as shown in the inset of B.
but also regional ones such as immigration from a
metapopulation (Shmida & Wilson 1985, Hubbell
2001). In the present case, the study area was cov-
ered by continuous forest, and even the most dis-
tant transects were not more than 25 km apart; so
it is reasonable to assume that the regional species
pool that was available to colonize a given site was
the same for all transects. Therefore, most of the
variability in species richness among transects was
likely to have depended on factors other than dif-
ferences in the regional species pool or immigration
rate, such as differences in local ecological condi-
tions.
The number of individuals per transect of both
plant groups was negatively correlated with soil
base content, but with species richness and diver-
sity, the trends differed. With Melastomataceae,
species richness and diversity also showed a nega-
tive correlation with soil base content, while with
pteridophytes, there was no correlation. This agrees
with the nding that the two plant groups showed
different trends in the distribution of species opti-
ma for soil base content (the weighted mean soil
base content of sites where each species was found;
Fig. 7). With pteridophytes, the optima were rel-
atively evenly distributed along the soil gradient,
although there was a tendency for intermediate val-
ues to be more common than extreme ones. In
contrast, most Melastomataceae had relatively low
optima for soil base content. Such different distri-
butions probably reect real differences in the eco-
logical preferences of the two plant groups.
Such contrasting trends in the relationship be-
tween species richness and soil base content be-
tween pteridophytes and Melastomataceae suggest
Pteridophytes and Melastomataceae in Yasun 531
that species richness of one group of organisms is
not necessarily a very good predictor of species
richness of another group of organisms. This result
agrees with earlier studies conducted both in the
tropics and elsewhere (Prendergast et al. 1993,
Howard et al. 1998). It may be difcult or even
impossible to nd a group of organisms that could
serve as a reliable indicator of spatial species rich-
ness patterns in other groups.
The soils in all of our Yasun transects have
relatively high base contents when compared to
other Ecuadorian and Peruvian sites that we have
inventoried. Therefore, the species richness trends
observed here represent only a small part of the soil
variation found in Amazonia, and the patterns ob-
served may not apply to longer edaphic gradients.
In earlier studies, we noticed that in soils with low-
er base contents, both pteridophytes and the Me-
lastomataceae showed a trend of increasing species
richness with increasing soil base content (Tuomis-
to & Ruokolainen 1994, Tuomisto & Poulsen
1996, Tuomisto 1998; K. Ruokolainen, pers. obs.).
These kinds of differences in species richness
trends in different parts of the edaphic gradient
probably explain some of the discrepancies among
the results of earlier studies. Several researchers
have suggested that the response curve of plant spe-
cies richness along a productivity gradient is uni-
modal and attains a peak at intermediate produc-
tivity (Ashton 1992, Rosenzweig & Abramsky
1993). If species richness along a long soil base
content gradient shows this kind of a unimodal
pattern, then the pattern observed along a short
edaphic gradient can be either decreasing, increas-
ing, or at depending on which part of the long
gradient is sampled (review in Pausas & Austin
2001).
In both pteridophytes and the Melastomata-
ceae, more than half of the common species
showed a statistically signicant bias in their abun-
dance distribution in relation to soil base content
(Figs. 2, 3). There is no reason to consider the soil
gradient observed here as exceptionally long in
western Amazonian terms, and therefore, our re-
sults suggest that the majority of pteridophyte and
Melastomataceae species perceive the unooded
forests of the area as a mosaic of different habitats,
rather than homogeneous. The fact that the species
optima were relatively evenly distributed along the
soil cation gradient, however, indicates that the
habitats need not be sharply differentiated from
each other either oristically or in terms of their
environment, but rather that they form a contin-
uum.
Patterns in pteridophyte and Melastomataceae
species composition have been observed to be high-
ly correlated with patterns in tree species compo-
sition (Tuomisto et al. 1995, Ruokolainen et al.
1997, Ruokolainen & Tuomisto 1998, Vormisto et
al. 2000). Therefore, one may expect that not only
pteridophytes and Melastomataceae but also many
tree species, and perhaps species of other plant
groups, show nonrandom distributions across an
edaphic gradient of the length observed in the pre-
sent study. Our results, however, cannot be extrap-
olated to other plant groups without further studies
because big plants may actually show more gener-
alized behavior along edaphic gradients than small
plants (Ruokolainen & Vormisto 2000, Ruokolai-
nen et al. 2002). The tree studies of Clark et al.
(1999) in Costa Rica and Webb and Peart (2000)
on Borneo support our conclusions, while those of
Pitman et al. (1999, 2001) in southern Peru sug-
gest that edaphic differences are less signicant.
Out of these studies, however, the Webb and Peart
(2000) study is the only one to include chemical
analyses of soils.
A comparison of species lists (Tables 1 and 2)
and soil properties for Yasun and other study sites
is illuminating. Out of the 49 pteridophyte species
considered rich-soil specialists by Tuomisto and
Poulsen (1996), 45 were found in our Yasun sam-
ple, while out of the 28 intermediate to poor soil
specialists, only 3 were. Similarly, only 4 out of the
24 Melastomataceae species reported by Tuomisto
and Ruokolainen (1994) from a poor to interme-
diate soil site in northern Peru were found in our
present Yasun sample, and all of these seem to be
generalists that have been found also elsewhere in
a wide range of soils (K. Ruokolainen, pers. obs.).
Such results suggest that the observed edaphic pref-
erences of pteridophyte and Melastomataceae spe-
cies are rather consistent even over wider spatial
scales.
ACKNOWLEDGMENTS
We thank Ponticia Universidad Catolica del Ecuador for
logistic support, especially its Yasun Research Station and
Herbarium QCA. Renato Valencia is acknowledged for
help in making the logistic arrangements, Oscar Mendoza
and Ermindo Zambrano for assistance in the eld work,
and Alan R. Smith and John T. Mickel for help with
species identication. The study was funded by the Dan-
ish Center for Tropical Biodiversity and the Academy of
Finland.
532 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Can as, and Celi
LITERATURE CITED
ASHTON, P. S. 1992. Species richness in plant communities. In P. L. Fiedler and S. K. Jain (Eds.). Conservation
Biology, pp. 322. Chapman and Hall, New York, New York.
BALSLEV, H., J. LUTEYN, B. LLGAARD, AND L. B. HOLM-NIELSEN. 1987. Composition and structure of adjacent
unooded and oodplain forest in Amazonian Ecuador. Opera Bot. 92: 3757.
CAMPBELL, D. G., D. D. DALY, G. T. PRANCE, AND U. N. MACIEL. 1986. Quantitative ecological inventory of terra
rme and varzea tropical forest on the Rio Xingu, Brazilian Amazon. Brittonia 38: 369393.
CAN

ADAS, L. 1983. El mapa bioclimatico ecologico del Ecuador. Editores Asociados C a. Ltda., Quito. Ecuador.
CLARK, D. B., D. A. CLARK, AND J. M. READ. 1998. Edaphic variation and the mesoscale distribution of tree species
in a neotropical rain forest. J. Ecology 86: 101112.
, M. W. PALMER, AND D. A. CLARK. 1999. Edaphic factors and the landscape-scale distributions of tropical
rain forest trees. Ecology 80: 26622675.
CLINEBELL, R. R., II, O. PHILLIPS, A. H. GENTRY, N. STARK, AND H. ZUURING. 1995. Prediction of neotropical tree
and liana species richness from soil and climatic data. Biodiv. Conserv. 4: 5690.
DUIVENVOORDEN, J. F. 1994. Vascular plant species counts in the rain forests of the middle Caqueta area, Colombian
Amazonia. Biodiv. Conserv. 3: 685715.
. 1995. Tree species composition and rain forest-environment relationships in the middle Caqueta area, Co-
lombia, NW Amazonia. Vegetatio 120: 91113.
. 1996. Patterns of tree species richness in rain forests of the middle Caqueta area, Colombia, NW Amazonia.
Biotropica 28: 142158.
DUQUE, A., M. SA

NCHEZ, J. CAVELIER, J. F. DUIVENVOORDEN, P. MIRAN

A, J. MIRAN

A, AND A. MATAPI

. 2001. Relacion
bosque-ambiente en el Medio Caqueta, Amazon a colombiana. In J. F. Duivenvoorden, H. Balslev, J. Cavelier,
C. Grandez, H. Tuomisto, and R. Valencia (Eds.). Evaluacion de recursos forestales no maderables en la
Amazon a noroccidental, pp. 99129. IBED, Universiteit van Amsterdam, Amsterdam, The Netherlands.
GENTRY, A. H. 1981. Distributional patterns and an additional species of the Passiora vitifolia complex: Amazonian
species diversity due to edaphically differentiated communities. Plant Syst. Evol. 137: 95105.
. 1988. Changes in plant community diversity and oristic composition on environmental and geographical
gradients. Ann. Mo. Bot. Gard. 75: 134.
GRANDEZ, C., A. GARCI

A, A. DUQUE, AND J. F. DUIVENVOORDEN. 2001. La composicion or stica de los bosques en

las cuencas de los r os Ampiyacu y Yaguasyacu (Amazon a peruana). In J. F. Duivenvoorden, H. Balslev, J.
Cavelier, C. Grandez, H. Tuomisto, and R. Valencia (Eds.). Evaluacion de recursos forestales no maderables
en la Amazon a noroccidental, pp. 163176. IBED, Universiteit van Amsterdam, Amsterdam, The Nether-
lands.
HOWARD, P. C., P. VISKANIC, T. R. B. DAVENPORT, F. W. KIGENYI, M. BALTZER, C. J. DICKINSON, J. S. LWANGA, R. A.
MATTHEWS, AND A. BALMFORD. 1998. Complementarity and the use of indicator groups for reserve selection
in Uganda. Nature 394: 472475.
HUBBELL, S. P. 2001. The unied neutral theory of biodiversity and biogeography. Monographs in population biology
32. Princeton University Press, Princeton, New Jersey.
HUSTON, M. 1980. Soil nutrients and tree species richness in Costa Rican forests. J. Biogeogr. 7: 147157.
JONGMAN, R. H. G., C. J. F TER BRAAK, AND O. F. R. VAN TONGEREN (EDS.). 1995. Data analysis in community and
landscape ecology. Cambridge University Press, Cambridge, United Kingdom.
PAUSAS, J. G., AND M. P. AUSTIN. 2001. Patterns of plant species richness in relation to different environments: An
appraisal. J. Veg. Sci. 12: 153166.
PITMAN, N. C. A., J. TERBORGH, M. R. SILMAN, AND P. NUN

EZ V. 1999. Tree species distributions in an upper

Amazonian forest. Ecology 80: 26512661.
, , , , D. A. NEILL, C. E. CERO

N, W. A. PALACIOS, AND M. AULESTIA. 2001. Dominance

and distribution of tree species in upper Amazonian terra rme forests. Ecology 82: 21012117.
POULSEN, A. D., AND H. BALSLEV. 1991. Abundance and cover of ground herbs in an Amazonian rain forest. J. Veg.
Sci. 2: 315322.
PRENDERGAST, J. R., R. M. QUINN, J. H. LAWTON, B. C. EVERSHAM, AND D. W. GIBBONS. 1993. Rare species, the
coincidence of diversity hotspots and conservation strategies. Nature 365: 335337.
ROMERO-SALTOS, H., R. VALENCIA, AND M. J. MACI

A. 2001. Patrones de diversidad, distribucion y rareza de plantas

lenosas en el Parque Nacional Yasun y la Reserva E

tnica Huaorani, Amazon a ecuatoriana. In J. F. Duiven-

voorden, H. Balslev, J. Cavelier, C. Grandez, H. Tuomisto, and R. Valencia (Eds.). Evaluacion de recursos
forestales no maderables en la Amazon a noroccidental, pp. 131162. IBED, Universiteit van Amsterdam,
Amsterdam, The Netherlands.
ROMOLEROUX, K., R. FOSTER, R. VALENCIA, R. CONDIT, H. BALSLEV, AND E. LOSOS. 1997. Arboles y arbustos (dap1
cm) encontrados en dos hectareas de un bosque de la Amazon a ecuatoriana. In R. Valencia and H. Balslev
(Eds.). Estudios sobre diversidad y ecolog a de plantas, pp. 189215. Memorias del II Congreso Ecuatoriano
de Botanica. Ponticia Universidad Catolica del Ecuador, Quito, Ecuador.
ROSENZWEIG, M. L., AND Z. ABRAMSKY. 1993. How are diversity and productivity related? In R. E. Ricklefs and D.
Schluter (Eds.). Species diversity in ecological communities, pp. 5265. The University of Chicago Press,
Chicago, Illinois.
RUOKOLAINEN, K., A. LINNA, AND H. TUOMISTO. 1997. Use of Melastomataceae and pteridophytes for revealing phy-
togeographic patterns in Amazonian rain forests. J. Trop. Ecol. 13: 243256.
Pteridophytes and Melastomataceae in Yasun 533
, AND H. TUOMISTO. 1998. Vegetacion natural de la zona de Iquitos. In R. Kalliola and S. Flores Paitan (Eds.).
Geoecolog a y desarrollo amazonico: estudio integrado en la zona de Iquitos, Peru. Ann. Univ. Turkuensis Ser.
A II 114: 253365.
, , J. VORMISTO, AND N. PITMAN. 2002. Two biases in estimating range sizes of Amazonian plant species.
J. Trop. Ecol. 18: 935942.
, AND J. VORMISTO. 2000. The most widespread Amazonian palms tend to be tall and habitat generalists. Basic
Appl. Ecol. 1: 97108.
SHMIDA, A., AND M. V. WILSON. 1985. Biological determinants of species diversity. J. Biogeogr. 12: 120.
SVENNING, J.-C. 1999. Microhabitat specialization in a species-rich palm community in Amazonian Ecuador. J. Ecol.
87: 5565.
TUOMISTO, H. 1998. What satellite imagery and large-scale eld studies can tell about biodiversity patterns in Ama-
zonian forests. Ann. Mo. Bot. Gard. 85: 4862.
, AND A. D. POULSEN. 1996. Inuence of edaphic specialization on pteridophyte distribution in neotropical
rain forests. J. Biogeogr. 23: 283293.
, AND . 2000. Pteridophyte diversity and species composition in four Amazonian rain forests. J. Veg.
Sci. 11: 383396.
, , AND R. C. MORAN. 1998. Edaphic distribution of some species of the fern genus Adiantum in
Western Amazonia. Biotropica 30: 392399.
, , K. RUOKOLAINEN, R. C. MORAN, C. QUINTANA, J. CELI, AND G. CAN

AS. Linking oristic patterns

with soil heterogeneity and satellite imagery in Ecuadorian Amazonia. Ecol. Appl. In press.
, AND K. RUOKOLAINEN. 1994. Distribution of Pteridophyta and Melastomataceae along an edaphic gradient
in an Amazonian rain forest. J. Veg. Sci. 5: 2534.
, , R. KALLIOLA, A. LINNA, W. DANJOY, AND Z. RODRIGUEZ. 1995. Dissecting Amazonian biodiversity.
Science 269: 6366.
VALENCIA R., R. CONDIT, K. ROMOLEROUX, G. VILLA, E. LOSOS, H. BALSLEV, J.-C. SVENNING, AND E. MAGAARD. Patterns
of tree species diversity and distribution from a large-scale forest inventory in Yasun National Park, eastern
Ecuador. In E. Losos and R. Condit (Eds.). The demography of tropical forest plants. Smithsonian Institution
Press, Washington, D.C. In press.
VAN REEUWIJK, L. P. 1993. Procedures for soil analysis. International Soil Reference and Information Centre. Technical
paper 9 (4th edition). Wageningen, The Netherlands.
VORMISTO, J., O. PHILLIPS, K. RUOKOLAINEN, H. TUOMISTO, AND R. VA

SQUEZ. 2000. A comparison of small-scale

distribution patterns of four plant groups in an Amazonian rainforest. Ecography 23: 349359.
WEBB, C. O., AND D. R. PEART. 2000. Habitat associations of trees and seedlings in a Bornean rain forest. J. Ecol.
88: 464478.
BIOTROPICA 35(1): 131 2003
ERRATUM
Biotropica 34(4)
IN TABLE 1 [Biotropica 34(4): 520521 by Tuomisto et al.], the data for D. falcata follow the collection
number 6259. The data following collection number 6064 belong to D. nodosa. The correct collection
number for D. nodosa is 6064 (not 6046).
131