Hydrobiologia (2015) 750:245–255
DOI 10.1007/s10750-014-2040-0
TRENDS IN AQUATIC ECOLOGY
The importance of forest cover for fish richness
and abundance on the Amazon floodplain
Javior Lobón-Cerviá • Laura L. Hess •
John M. Melack • Carlos A. R. M. Araujo-Lima
Received: 6 June 2014 / Revised: 30 August 2014 / Accepted: 6 September 2014 / Published online: 25 September 2014
Ó Springer International Publishing Switzerland 2014
Abstract Flooded forest is one of the most important
fish habitats in Neotropical rivers, and one that is
increasingly subjected to negative impacts from
logging, agriculture, and other human activities. The
purpose of our study was to test quantitatively whether
fish richness and abundance in Amazonian floodplain
lakes are associated with the area of flooded forest. We
sampled fish and several other variables in 35 Ama-
zonian floodplain lakes during the high-water season.
Our results highlighted that fish richness and abun-
dance were directly related to flooded forest, inversely
related to distance from the river, and influenced by
dissolved oxygen concentration\1 mg l-1. The same
result applied to fish richness and abundance landed by
fisheries. Other variables such as depth and area of
Carlos A. R. M. Araujo-Lima—deceased.
Guest editor: Koen Martens / Emerging Trends in Aquatic
Ecology
J. Lobón-Cerviá
Museo Nacional de Ciencias Naturales (CSIC), C/ Jose
Gutierrez Abascal, 2, 28006 Madrid, Spain
L. L. Hess
J. M. Melack (&)
Earth Research Institute, University of California,
Santa Barbara 93106, USA
e-mail: melack@bren.ucsb.edu
C. A. R. M. Araujo-Lima
Instituto Nacional de Pesquisas da Amazônia, Manaus,
Brazil
open water habitat were also related but the results
were less consistent and apparently reliant on sam-
pling methodology. Our results suggest that conser-
vation of the flooded forest is critical for the
maintenance of fish assemblages in the Amazon, and
that removal of flooded forest will reduce fish richness,
fish abundance, and fisheries yield.
Keywords Amazon
Flooded forests
Fishes

Fisheries
Synthetic aperture radar
Introduction
Fish have an important socio-economic role for human
communities living along tropical rivers and are a
major protein source for these people (Daget, 1973;
Fabré & Alonso, 1998; Cerdeira & Ruffino, 2000;
Santos & Santos, 2005; Batista & Issac, 2012; Santos
et al., 2014). Because the annual flood reduces the area
available to agriculture, local farmers turn to fisheries
to obtain income during the high-water season. Over
the past 30 years, pressure on floodplain fish stocks in
the Amazon basin has increased owing to changes in
fishing technology, increased demand for fish pro-
ducts, a shift in the local economy from farming to
fishing, and the expansion of cattle ranching on the
floodplain, with accompanying deforestation (McG-
rath et al., 1999; Castello et al., 2013). Over the past
150 years, boom and bust cycles of rubber, cacao, and
123
246
jute production have been associated with deforesta-
tion on the mainstem Amazon floodplain downstream
of Manaus (WinklerPrins, 2006), and agriculture and
livestock activities continue to be important drivers of
floodplain deforestation (Sheikh et al., 2006; Renó
et al., 2011). Negative impacts on fish habitat of both
large hydroelectric projects and smaller stream
impoundments are of increasing concern (Finer &
Jenkins, 2012; Macedo et al., 2013). Recent severe
droughts provide raise further issues for floodplain
habitats (Tomasella et al., 2013).
Riparian forests are an important habitat for
freshwater fish worldwide, because forests provide
fish with both shelter and food including fruits, seeds,
and canopy insects (Goulding, 1980; Lowe-McCon-
nell, 1987; Gerking, 1994). Seasonally inundated
riparian forests, often called flooded forest or varzea
in the Amazon basin, are extensive in tropical South
America and Africa, where they form one of the main
habitats on the floodplains (Mayaux et al., 2002; Junk
et al., 2010; Melack & Hess, 2010).
Despite ongoing deforestation in tropical areas,
little information exists concerning its impact on
tropical fish and fisheries. Several studies have
addressed the relation between forest cover and stream
fishes in tropical areas (Burcham, 1988; Lyons et al.,
1995; Toham & Teugels, 1999; Bojsen & Barriga,
2002). Similar studies have not been done for large
rivers, even though destruction of habitats supporting
fish populations has been hypothesized to be an
important factor affecting the long-term health of
fisheries.
Studies of the relationship between forest cover and
stream fish have reached inconsistent conclusions. In
Costa Rica, species richness was found to be higher in
pasture streams than in forest streams (Burcham,
1988), but such differences were not found in Mexico
(Lyons et al., 1995) or Ecuador (Bojsen & Barriga,
2002). In addition, Toham & Teugels (1999) and
Bojsen & Barriga (2002) found greater fish abundance
in forest-covered streams of Mexico and Ecuador than
in deforested streams, but Lyons et al. (1995) reported
inverse relationships for streams in Mexico. A possi-
ble explanation for the inverse relationship between
fish abundance and richness and the presence of
riparian forest is the higher availability of autochtho-
nous food in deforested streams. Because open
streams receive more light and therefore have higher
algae growth than shaded streams, they tend to have a
123
Hydrobiologia (2015) 750:245–255
higher density of algivorous fishes (Powers, 1984).
Algivores and detritivores are abundant in Neotropical
rivers, and increasing primary production may allow
more species to coexist.
Efforts to evaluate relationships between attributes
of fish assemblages and area of flooded forest have
been limited by the difficulty of measuring the
vegetation cover of inundated forest in the humid
tropics. Synthetic aperture radar (SAR) imagery
alleviates this limitation. This tool can recognize
forests and other types of vegetation and reveal water
beneath the canopy, allowing reasonable estimation of
the amount of flooded forest, aquatic herbaceous
plants, and open water habitats in floodplains (Hess
et al., 1995, 2003). However, habitat availability is
only one factor that may affect fish richness and
abundance in floodplains. Junk et al. (1983) reported
that assemblage structure changed when dissolved
oxygen availability was reduced. Relationships among
depth, piscivores, and fish abundance have been
reported in a few studies (Winemiller et al., 2000;
Súarez et al., 2001). Fisheries may also reduce the
abundance of fish locally. Periodicity or variability of
inundation can influence fish species richness (Jardine
et al., in press) and influence the structure of fish
communities (Sousa & Freitas, 2008), while high
water levels connect habitats leading to similarity in
biological communities (Thomaz et al., 2007).
The purpose of our study was to test whether fish
richness (i.e., number of species) and abundance are
associated with the availability of flooded forest
during high-water season on the Amazon floodplain.
The local importance of fish, the availability of SAR
mosaics obtained at high water, and the extent and
variability of flooded forest cover make the Amazon
basin an appropriate region in which to evaluate this
association. Fish abundance was analyzed as all
species, as those landed in the markets, and as
algivore–detritivore species. Our results provide evi-
dence for assessing the potentially deleterious effect of
floodplain deforestation on tropical fish and fisheries.
Methods
The seasonal flood is one of the most important
modulators of the Amazon landscape (Junk, 1997). In
the central Amazon, the water level on the floodplain
varies approximately 8–12 m annually. During the
Hydrobiologia (2015) 750:245–255
Fig. 1 Geographical
locations of the sampling
sites along the floodplain of
the central Solimões/
Amazon River. Numbers 1 2º S
3 4
refer to the lakes in Table 1 7 1
1 2526
2 5 8
6 27 35 31 24 28
9 34
10
high-water period, most lakes are connected via
channels to rivers and often to other lakes through
the floodplain. The three main types of habitats in
floodplain lakes are herbaceous vegetation (aquatic
macrophytes), riparian forest, and open water. Ripar-
ian forests border the lakes, and herbaceous plants are
usually located between the riparian forest or shore
and the open water. The relative proportions of the
three habitats vary regionally and seasonally (Hess
et al., 2003). Further descriptions of Amazon flood-
plain lakes, their ecology and ecosystem processes are
provided by Junk (1997), Melack & Forsberg (2001),
and Melack et al. (2009).
Classified mosaics of JERS-1 (Japanese Earth
Resources Satellite-1) SAR images were used for the
selection of sampling sites and calculation of habitat
areas. The L-band SAR images were acquired during
September and October 1995 (low-water stage) and
during May and June 1996 (high-water stage), and
mosaicked to create geo-referenced digital datasets
with a pixel size of 3 arc-seconds, approximately 93 m
(Chapman et al., 2002). After masking out upland
areas not subject to inundation, the mosaics were
classified into habitat types based on the radar
backscattering coefficient. The classified image was
field-checked in May 2000 in 12 areas with four or
more contiguous pixels of one habitat type, and found
to be 75% accurate. Further details of SAR processing,
classification, and validation are provided by Hess
et al. (2002, 2003).
Habitats present at high-water stage were flooded
forest, non-flooded forest (on high levees), aquatic
macrophytes, and open water. Riparian forests were
densely covered by tall trees (height [ 20 m) with
water under the canopy, including many fruit-
247
64º W 60º W
Manaus 29
30 36
33
32
19 17 18
20 23 13 14
12 11
21 22 15 16 4º S
producing species, and in some areas, emergent, small
trees such as Cecropia latiloba and Vitex cymosa.
Macrophytes were mostly floating grasses.
Thirty-five lakes were selected for sampling along
1300 km of the Amazon River floodplain (Fig. 1;
Table 1). Lakes were selected to include a range of
sizes and types as well as a range of riparian forest
cover. The lakes selected could be broadly classified
into three main types (Sippel et al., 1992): island lakes
(12 lakes), round lakes (7 lakes), and dendritic lakes
(16 lakes). Dendritic lakes included those with shapes
partly dendritic and partly round. Lake areas varied
from 0.01 to 18.9 km2, falling within the range of the
most common lake sizes on the Amazon floodplain
(Sippel et al., 1992).
Sampling site locations were measured with geo-
graphic positioning system (GPS) units in the field and
located on the classified SAR mosaics. Habitat areas
were measured on classified mosaics within buffers
with radii of 1.9, 3.9, and 7.8 km (with corresponding
areas of 11.3, 47.8, and 191 km2), centered on the
fishing sites. The buffers will be referred to as the 2-,
4-, and 8-km buffers. These radii were based on
estimates of maximum distance traveled by fish from
sampling sites, calculated using a mean fish body
length of 18 cm, a swimming speed of 1 body
length s-1, and periods of activity of 3, 6, and 12 h.
Buffer limits were manually adjusted to avoid the
main river channel and side channels. The area of
habitat types in each buffer was calculated after
correcting to an equal-area projection.
At each fishing site, dissolved oxygen concentra-
tion (DO, mg l-1), water temperature (°C), and
electrical conductivity (lS cm-1) were measured
in situ with a polarographic oxygen electrode,
123
 Table 1 Names, locations, and attributes of the 35 sampled lakes; numbers in brackets refer to the geographical location in Fig. 1
248

Name Lat Long Depth Cond DO T Houses D Area Ft Ft Ft W W W M M M

(°S) (°W) (m) (lS (mg l-1) (°C) (n) (km) (km2) 2 km 4 km 8 km 2 km 4 km 8 km 2 km 4 km 8 km

123
cm-1) (km2) (km2) (km2) (km2) (km2) (km2) (km2) (km2) (km2)

Unknown (1) 3.8953 62.5930 12.0 96 2.69 27.6 1 0.8 0.27 3.4 9.9 13.2 1.2 1.6 1.9 2.1 3.5 4.1
Ipixuna (2) 3.8553 63.8797 10.4 75 2.14 26.2 300 1.8 3.46 2.0 8.8 23.9 2.8 4.6 5.2 0.7 2.9 5.5
Cipotuba (3) 3.8140 62.3451 16.0 93 2.28 28.0 1 1.5 1.86 6.5 25.1 58.2 1.6 2.8 8.6 3.3 3.8 9.8
Anori (4) 3.7725 61.6997 9.8 78 0.53 26.6 1495 7.6 0.90 8.7 31.1 128.7 1.2 7.1 19.6 2.6 6.0 20.6
U.Grande (5) 3.9235 62.0302 12.0 90 3.15 28.3 8 3.1 6.83 5.3 26.8 103.5 4.8 10.0 14.8 2.4 3.9 9.3
S. Tomé (6) 3.9398 61.4812 11.5 90 0.68 27.3 6 2.6 3.27 4.6 23.2 82.0 4.8 7.8 8.3 3.1 8.9 29.7
Rasgado (7) 3.6060 61.1632 8.0 84 2.51 27.2 95 1.3 0.84 3.1 12.4 21.2 1.4 2.3 7.7 4.7 5.8 3.4
Jacaré (8) 3.6568 60.8067 9.5 39 1.23 27.8 45 6.3 1.36 3.4 18.1 86.3 2.4 7.9 12.3 3.5 8.6 16.3
Acurú (9) 3.6025 63.6795 12.0 29 1.33 27.0 52 4.3 0.00 3.0 22.8 85.3 1.1 2.2 77.4 0.6 3.2 5.1
Comprido (10) 3.7488 63.5458 11.4 68 0.49 26.7 1 4.9 2.35 4.7 27.8 126.1 3.8 10.8 23.1 4.0 6.8 18.3
Apaurá (11) 3.8987 63.4327 8.1 93 3.81 27.6 72 0.6 5.46 1.5 8.5 22.0 4.2 6.5 11.3 2.1 3.0 4.6
Ajurá (12) 3.9544 62.7659 9.4 87 1.67 27.8 3 1.9 1.55 4.5 15.4 42.3 1.1 2.3 3.3 0.6 1.6 3.0
Luis (13) 3.9073 62.8017 10.0 99 0.31 26.8 10 1.0 1.37 6.9 15.2 33.4 1.0 2.7 4.2 1.1 1.4 2.4
Tarará (14) 2.4225 66.5028 10.0 120 2.86 25.8 15 0.4 0.73 3.4 15.1 35.6 0.6 1.5 3.0 2.7 3.5 6.8
Campina (15) 2.6139 66.3472 7.0 91 0.22 25.6 1 1.8 0.05 4.6 23.2 65.5 1.8 2.5 4.9 6.1 9.2 18.0
Curimatá (16) 2.0378 66.3339 7.0 88 2.47 27.4 120 2.0 3.18 1.4 8.7 50.2 8.8 17.7 9.4 2.0 5.6 8.6
Mirita (17) 1.9794 66.0422 6.0 70 1.49 26.9 28 2.8 0.55 2.4 20.4 59.2 3.0 3.3 5.0 3.1 3.1 6.2
Sacambu (18) 2.5671 65.8360 7.0 48 1.37 26.8 1 3.7 2.08 3.4 29.2 93.5 6.0 8.0 12.6 3.1 9.3 18.9
Tamaniquá (19) 2.7311 65.7335 10.0 71 2.28 26.4 40 3.6 2.03 6.2 35.4 133.0 1.5 2.2 8.3 2.4 3.0 8.8
Malvado (20) 2.3311 66.3550 7.0 124 2.68 26.9 6 9.5 0.65 5.8 38.9 152.4 0.2 1.4 6.9 3.4 4.9 21.7
Uará (21) 2.7110 65.6036 8.0 25 1.94 26.9 42 4.8 0.90 3.7 17.1 53.0 1.1 1.7 2.9 0.5 1.9 4.4
Alvaraes (22) 3.2052 64.8340 11.0 78 3.57 27.5 1333 2.8 0.82 3.3 10.4 20.0 1.3 1.3 1.7 0.5 1.5 2.3
Estasio (23) 2.8083 57.2044 8.0 239 1.64 27.6 8 1.5 3.92 1.2 10.3 23.9 0.2 7.1 11.9 0.3 7.7 17.6
Cararaçú (24) 2.3646 57.5990 8.5 21 3.00 26.3 1 5.4 0.95 0.6 1.3 3.2 1.2 2.2 9.2 5.0 5.8 18.4
Terra preta (25) 2.4442 57.6381 8.0 75 1.40 27.6 1 4.8 5.52 0.1 0.3 7.3 2.8 6.1 9.8 2.9 9.0 15.8
Apunumá (26) 2.9149 58.1130 8.0 31 2.80 27.9 1 0.6 1.46 2.5 5.6 7.7 2.6 3.7 3.9 4.2 3.5 3.5
Piloto (27) 2.7798 57.0921 6.3 56 2.80 28.5 4 0.9 7.35 0.8 8.7 22.8 7.2 11.0 38.5 3.0 5.6 20.4
Arari (28) 2.5111 57.1956 6.5 47 1.91 27.8 2 1.4 8.74 3.3 5.1 24.3 0.2 10.3 12.9 9.0 10.6 17.2
Corócoró (29) 3.2276 58.6921 9.0 25 0.67 27.9 1 5.4 0.10 2.4 37.6 114.7 0.5 1.1 9.5 3.9 4.1 19.5
Moaná (30) 2.8690 57.5258 8.0 206 4.20 28.8 75 1.4 10.87 5.2 8.7 24.2 2.5 13.3 16.1 4.9 7.4 14.6
Taperebá (31) 3.3117 58.6618 10.2 25 1.62 28.2 10 2.4 5.09 3.0 18.6 40.9 2.6 7.1 12.4 5.4 13.2 27.6
Garças (32) 3.1795 57.6967 7.3 92 4.54 29.2 50 4.2 3.65 0.7 24.1 57.0 0.0 4.1 6.1 0.0 4.5 9.9
Paracuubá (33) 3.0644 59.2522 12.0 12 2.35 27.7 50 10.5 0.13 1.1 2.1 7.0 0.1 0.6 7.5 1.2 2.4 20.0
Hydrobiologia (2015) 750:245–255
Hydrobiologia (2015) 750:245–255 249

Cond electrical conductance, DO dissolved oxygen, T water temperature, Houses number of houses, D distance from river, Ft, W, and M the habitats flooded forest (Ft), open water (W), and macrophytes
thermistor, and conductivity sensor, respectively. DO

(km2)
8 km

26.7
21.0
was measured in the water column every meter. The
M

maximum depth (m) of lakes was determined by sonar.

Households around the lake were counted directly or
(km2)
4 km

8.9
9.8 by interviewing community leaders. Households were
M

used as an index of potential fishing pressure. Lake

area (km2) during low water and distance from the
(km2)
2 km

5.0
4.0
M

main river channel (km) were measured on the

mosaics.
(km2)
8 km

16.4
17.7

Fish were sampled with a set of 12 gill nets with

W

3–16 cm mesh sizes (between opposite knots). All

nets were 25 m long and 2.5 m high, except those with
(km2)
4 km

5.9
10.7

3 and 16 cm mesh, which were 1.9 and 3.2 m high,

W

respectively. A new set of nets was used per lake. To

(km2)
2 km

reduce seasonal variation of fish assemblages, we used

5.4
5.4
W

three independent boats and teams, each one sampling

one section of the river basin (i) 57.1 W–59.0 W; (ii)
(km2)
8 km

63.1
16.5

60.8 W–63.8 W; and (iii) 64.8 W–66.5 W. The sam-

Ft

pling season was chosen to capture the near maximum

species richness (Mérona & Bittencourt, 1993) and
(km2)
4 km

25.4
8.7

stability in water level. Due to a small longitudinal

Ft

flood lag, we started sampling in the west side of the

(km2)

basin.
2 km

2.1
2.1
Ft

Gill nets were set at the edge of the flooded forest at

(km2)

3.1 m depth [standard deviation (SD), 0.7], to reduce

Area

2.74
8.11

dolphin attacks and improve their performance. We

inspected the nets and measured DO and temperature
(km)

2.9
3.0
D

every 4 h during 24 h. Fish were packed after fixation

Houses

with 10% formalin. Fish identification, measuring and

(n)

weighing were conducted in the laboratory at the

2
5

Instituto Nacional de Pesquisas da Amazônia. Species

27.4
28.7
(°C)

landed by the commercial fleet were classified

T

following Ferreira et al. (1998). Catch was expressed

(mg l-1)

as 1000 m-2 day-1 (Catch Per Unit Effort, CPUE)

1.84
3.12
DO

and kg 1000 m-2 day-1 (Biomass Per Unit Effort,

cm-1)

BPUE) to allow comparison with other studies and to

Cond

38
113
(lS

correct the data from three lakes, where a few nets

were lost. CPUE and BPUE include juveniles and
(M) within the three buffers of 2, 4, and 8 km
Depth
(m)

7.0
5.2

adults.
We tested the effect of several independent vari-
57.9962
57.7464
Long
(°W)

ables on the species richness, CPUE, and BPUE of all

species, commercially important species, and detriti-
2.7775
3.0619

vores with a general linear model. Because we could

(°S)
Lat

not predict which buffer size would relate better to the

Table 1 continued

dependent variables, we built three models, one for

each buffer size. One categorical (dummy) variable
Buiuçú (35)
Arroz (34)

was added to the initial models: critical mean DO. This

variable was set to 1 when mean DO was above
Name

1 mg l-1 and to 0 when DO was lower than 1 mg l-1.

123
250
This DO value is the critical concentration for many
hypoxia sensitive species in the Amazon (Val &
Almeida-Val, 1995). Heteroscedasticity was corrected
by log-transforming the dependent variables. To
develop the statistical model, we first used the partial
residuals to evaluate the fitness of variables to the
model, and then fit the variables previously selected to
the model.
Results
The maximum depth of the lakes sampled ranged from
5.2 to 16 m; their areas reduced during the low-water
season, but only a few lakes dried completely.
Electrical conductivity ranged from 12 to
239 lS cm-1, indicating a mixture of river and upland
runoff in the more dilute waters (Table 1). The lakes
have variable human populations (0–1445 house-
holds) on their periphery.
Most sampling sites (70%) were located within
4 km of the Amazon River main stem. Daily mean and
minimum DO of the water column varied from 0.2 to
4.5 and 0.0 to 2.7 mg l-1, respectively. In six lakes,
the mean DO was lower than the critical value
(1 mg l-1). Mean temperature of water columns
ranged from 25.6 to 29.2°C. The relative area of
habitats changed with the size of the buffer zones.
Flooded forest was the largest habitat in the 4 and
8 km buffers, followed by herbaceous vegetation and
open water. However, all habitats were equally
extensive in the 2 km buffer. We collected 8122 fish
belonging to 195 species (species list in Granado-
Lorencio et al., 2005; further information in Granado-
Lorencio et al., 2007). The average and range of
CPUE, BPUE, and species richness at sampling sites
are given in Table 2.
The first step of the statistical analyses was to
evaluate the correlation between the independent
variables, which could lead to co-linearity. We found
that in the 4- and 8-km buffers there were significant
correlations between three variables: longitude, tem-
perature, and area of the lake during low-water season.
We excluded temperature and longitude from the
analyses because they seemed less related to the
dependent variables. Second, another exploratory
analysis showed significant interactions between crit-
ical DO and flooded forest area, open water area, and
distance to the river. Therefore, as a first adjustment,
123
Hydrobiologia (2015) 750:245–255
Table 2 Mean, SD and range of species richness, catch per
unit of effort in number and biomass for all species (t), landed
species (com), and detritivores (d) of 35 Amazonian floodplain
lakes
Mean SD Range
Species richness 41 13 15–80
CPUEt (fish/1000 m -2 -1
day ) 298 153 81–681
BPUEt (kg/1000 m-2 day-1) 50 26 7–105
CPUEcom (fish/1000 m-2 day-1) 248 134 69–562
BPUEcom (kg/1000 m-2 day-1) 45 24 5–92
CPUEd (fish/1000 m-2 day-1) 69 91 1–367
BPUEd (kg/1000 m-2 day-1) 9 10 0.1–41
we split the data into lakes with DO above and below
the critical level. The number of lakes with mean DO
below the critical level was six, too few to allow model
fitting.
With the dataset reduced to 29 observations, we
tested the coefficient of the independent variable to
evaluate which explained the variability in species
richness and CPUE (Table 3). Two variables were
consistently related to species richness and CPUE of
total and of commercial species: distance from the
river and area of flooded forest. Other variables such as
depth, average DO, and open water area were signif-
icant in some of the regressions. Conductivity, number
of households, temperature, area of the lake during
low-water season, and the area of aquatic macrophytes
were not significant in any of the multiple regressions.
The exception was CPUE and BPUE for detritivores;
regressions included only the distance from the river
and DO.
The species richness, abundance, and biomass of all
fishes and of commercially important fish increased
significantly with the amount of flooded forest in the
three buffer sizes and decreased with the distance of
the sampling site from the main river channel
(Table 4). Total CPUE decreased with average DO
and increased with the area of open water habitat and
with maximum depth in some buffer sizes. In most
models, the independent variables explained at least
50% of the variance in fish abundance, reaching nearly
80% in a few cases. The CPUE and BPUE of
detritivores were inversely related to the distance
from the main river and with DO.
In all cases, the residuals showed no consistent
tendencies. Durbin-Watson statistics were not signif-
icantly different than 2 and revealed no residual
Hydrobiologia (2015) 750:245–255 251

Table 3 Coefficients of multiple regressions for species richness and catch per unit of effort in number and biomass for total species
(t), landed species (com), and detritivores (d) versus seven environmental variables within three buffers
Variable Buffer Depth Cond DO T Houses Area D (km) Forest Water Macr
(km) (m) (lS cm-1) (mg l-1) (°C) (n) (km2) (km2) (km2) (km2)

Richness 2 0.03 0.37 0.53 0.03 0.11 0.31 2.94* 1.72 1.24 0.36
Richness 4 1.13 0.38 1.57 0.69 1.57 2.77* 2.10* 0.83 0.79
Richness 8 1.59 0.34 0.87 0.13 0.62 2.29* 1.96* 0.07 0.19
lnCPUEt 2 1.57 0.43 2.77* 0.35 0.05 1.29 3.19** 2.90** 3.07** 0.92
lnCPUEt 4 3.23** 0.21 2.57* 0.30 0.86 4.31*** 4.09*** 3.03** 0.24
lnCPUEt 8 2.45* 0.42 2.41* 0.01 1.68 3.88** 3.76** 1.03 0.39
lnBPUEt 2 0.30 0.39 0.39 1.18 1.17 0.22 3.15** 1.27 1.20 0.76
lnBPUEt 4 1.06 0.40 1.38 0.57 0.36 3.61** 2.77* 1.06 0.49
lnBPUEt 8 0.80 0.84 1.38 0.14 0.93 3.09** 3.14** 0.71** 1.29
lnCPUEc 2 2.11* 0.63 3.21** 0.56 0.23 1.01 3.64** 2.74* 4.51*** 0.25
lnCPUEc 4 2.75* 0.13 2.01 0.54 0.26 4.021** 2.59* 3.31** 0.49
lnCPUEc 8 1.92 0.11 2.20* 0.27 1.61 3.73** 2.89** 1.28 0.20
lnBPUEc 2 0.67 1.52 1.62 0.24 1.27 0.58 3.36** 2.57* 2.18* 1.29
lnBPUEc 4 1.12 0.82 1.14 0.25 0.42 3.49** 2.34* 1.02 0.58
lnBPUEc 8 0.81 0.84 1.17 0.20 1.06 2.80* 2.51* 0.79 1.25
lnCPUEd 2 1.32 0.04 2.98** 0.88 0.77 1.96 2.09* 0.25 0.80 1.48
lnCPUEd 4 1.40 0.61 1.78 0.56 0.05 2.70* 0.68 1.15 0.32
lnCPUEd 8 1.10 0.86 2.06* 0.46 1.20 2.18* 1.08 0.46 0.75
lnBPUEd 2 1.14 1.38 0.16 1.66 0.47 1.48 1.39 1.08 0.11 1.15
lnBPUEd 4 0.45 1.40 1.21 0.37 0.25 2.87* 0.52 0.45 0.71
lnBPUEd 8 0.35 1.68 1.23 1.01 0.88 2.12* 0.82 0.42 1.16
Number of observations = 29
Cond electrical conductance, DO dissolved oxygen, D distance from river, Macr macrophytes
Significant coefficients are indicated as * 0.05 \ P \ 0.01; ** 0.009 \ P \ 0.001; *** P \ 0.0001. Values of temperature (T) are
excluded from the analyses in buffers 4 and 8 km to avoid collinearity

correlations; autocorrelation was below 0.2. Toler-
ances were always above 0.7, suggesting that no multi-
collinearities were occurring between independent
variables.
Discussion
Fish richness and abundance in the floodplain of the
Amazon River were associated with the amount of
flooded forest in the buffer areas. Four- and 8-km
buffers included most of the lakes and the associated
floodplain. Hence, we suggest that lakes with more
flooded forest have higher fish abundance, either in
number or biomass, than lakes with less flooded forest.
Other important factors were the proximity of the
sampling sites to the main river and the critical oxygen
level (\1 mg l-1). These results apply also to the
abundance of the fish species exploited by fisheries,
suggesting a real dependence of the local fisheries on
the flooded forest. Conversely, the abundance of
detritivorous fish was not associated with the amount
of flooded forest, but only with proximity to the main
river and DO.
Three variables were expected a priori to correlate
with the amount of fish caught in the floodplain, but
did not: number of people using the resources in the
lakes, lake area during low water, and the amount of
aquatic macrophytes. Residents on the riverbanks fish
intensively during the high-water season. They con-
sume large amounts of fish (300–500 g day-1) and
also sell part of the yield to earn income (Cerdeira
et al., 1997; Fabré & Alonso, 1998); therefore, lakes
with dense population should have fewer fish.

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252 Hydrobiologia (2015) 750:245–255

Table 4 Results of multiple linear regressions between species richness and catch per unit of effort in number and biomass for total
(t), landed (com), and detritivorous (d) fish versus nine environmental variables using three buffer sizes
Variable Buffer (km) Forest (km2) Water (km2) Dist. (km) DO (mg l-1) Depth (m) P r2

Richness 2 3.788*** NS -2.413** NS NS \0.001 0.50

Richness 4 0.483 NS -2.898 NS NS 0.002 0.38
Richness 8 0.14 NS -3.276 NS NS 0.002 0.39
lnCPUEt 2 0.140*** 0.080* -0.089*** -0.180** NS \0.001 0.70
lnCPUEt 4 0.022*** 0.054*** -0.106*** -0.176** 0.078** \0.001 0.79
lnCPUEt 8 0.007*** NS -0.149*** -0.154** NS \0.001 0.65
lnBPUEt 2 0.164*** NS -0.125*** NS NS \0.001 0.57
lnBPUEt 4 0.026** NS -0.151*** NS NS \0.001 0.53
lnBPUEt 8 0.009*** NS -0.177*** NS NS \0.001 0.61
lnCPUEc 2 0.137*** 0.104** -0.096*** -0.210** NS \0.001 0.78
lnCPUEc 4 0.020** 0.060** -0.102*** NS 0.077** \0.001 0.66
lnCPUEc 8 0.006** NS -0.157*** -0.194 NS \0.001 0.63
lnBPUEc 2 0.166** NS -0.135** NS NS \0.001 0.58
lnBPUEc 4 0.025** NS -0.16*** NS NS \0.001 0.51
lnBPUEc 8 0.008** NS -0.183*** NS NS \0.001 0.57
lnCPUEd 2–8 NS NS -0.286 -0.821 NS 0.003 0.35
lnBPUEd 2–8 NS NS -0.259 NS NS 0.023 0.18
Only significant independent variables in at least one regression are listed. Number of observations = 29; asterisks indicate the
probability of coefficients being equal to 0; *** P \ 0.001; ** 0.01 \ P [ 0.001; no asterisks = 0.05 \ P [ 0.01

Floodplain lakes are seasonal and water volume is
reduced during the dry season (Melack & Forsberg,
2001). If we consider that lake area approximates
relative lake volume, one would expect that lakes
which vary less in area would be more favorable
habitats than those where the area is reduced more
during the dry season, and that habitat stability favors
high species richness and abundance. This effect has
been reported in oxbow lakes of the Brazos River
(Winemiller et al., 2000) and, more generally, in a
comparison of Neotropical and Australia rivers (Jar-
dine et al., in press).
Rodriguez & Lewis (1994), Bayley (1989), and
Súarez et al. (2001) suggested that floodplain macro-
phytes and the abundance of piscivores are the main
factors structuring fish communities of the Amazon,
Pantanal, and Orinoco. Neither factor was correlated
with fish abundance in our study. Lake depth has often
been assigned as an important factor to explain fish
abundance. Winemiller et al. (2000) found a direct
relation between these two variables in oxbow lakes of
the Brazos River (USA), and Súarez et al. (2001) and
Tejerina-Garro et al. (1998) reported depth as one of
the main environmental structuring factors of fish
assemblages in the Pantanal and Araguaia River
(Brazil). Lake depth and open area had a weak effect
on the abundance of fish in our study; its correlation
was inconsistent and was dependent on buffer size.
The lack of relationship between these variables
and fish catch in the floodplains could be accounted for
by sampling limitations (type II error) or could
represent a real lack of effect. Since we studied 35
lakes, our statistical resolution was reasonable despite
the variability in the fish catch, suggesting that the
more probable hypothesis is that these variables had a
weak effect, if any, on fish richness and abundance.
Junk et al. (1983) and Val & Almeida-Val (1995)
argued that fish distribution is influenced by DO in
Amazonian floodplain lakes; our results agree. Lakes
with low DO (\1 mg l-1) did not fit well to the
regression models. However, the inverse relationship
between DO and fish abundance in lakes with
intermediate DO was unexpected: we cannot explain
this correlation, but it suggests that fish prefer sites
with median DO.
The most consistent variable, which, together with
the amount of flooded forest, predicted the fish catch,
was the distance of the sampling site from the river:

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Hydrobiologia (2015) 750:245–255
species richness and abundance are greater near the
river. Migratory species, which form an important
fraction of the assemblage, may prefer to disperse near
the river along their major migration route. Possibly,
the area near the river is an ecotone having channel
benthic species in addition to the floodplain species.
Fish richness and abundance consistently increased
with the amount of flooded forest surrounding the lake.
The forest provides food for fish (Goulding et al.,
1988) and this increases the overall food available for
consumers. In addition, the higher fish abundance in
the forest attracts piscivores, which may find more
prey in this complex habitat.
However, not all fish seem to depend on the forest.
The detritivores were not directly related to this
habitat, suggesting that they can find suitable detritus
in other habitats of the floodplain. This may conflict
with the assertion in Goulding et al. (1988) that
dissolved organic carbon produced by the flooded
forest is one of the main sources for detritus. Algae
exudates and particulate organic matter exported by
the streams probably are more important for this
trophic guild, as previously suggested by Araujo-Lima
et al. (1986). In addition, even for fish associated with
the flooded forest, other habitats may play a role
during a part of the life cycle; for example, floating
meadows provide shelter for juveniles (Henderson &
Robertson, 1999). In a study based on fish collections
over 11 months on a seasonally inundated island in the
Solimões River with extensive deforestation, Corredor
(2004) found that the fish community structure was
similar in habitats with open water, flooded forests,
and floating macrophytes. Based on sampling another
inundated island in the Solimões River, Petry et al.
(2003) reported that species richness was significantly
higher in vegetated areas than in unvegetated areas,
and stands of floating grasses had highest richness.
Both Melack et al. (2009) and Monteiro (2011)
reported weak and complex relations between flooded
areas and fish yields.
The dependence of fish on the flooded forest
elucidated in this study enhances the importance of
this habitat. What in the past was mostly a concern of
conservationists becomes a strategic resource for the
whole human society of the Amazon, which is highly
dependent on local fisheries. The consumption of fish
in the region is among the highest in the world
(Cerdeira et al., 1997; Fabré & Alonso, 1998), and
fisheries employ about 15% of the local populations.
253
As a consequence, any reduction of the flooded forest
for agriculture or cattle ranching will have an effect on
the number and abundance of species, including those
exploited by fisheries. For example, the results of this
study predict that a reduction of 50% of the flooded
forest of a lake, distant 1 km from the river, with
average DO equal to 2 mg l-1, average depth of 12 m,
and 5 km2 of open water, would cause a decrease of
25% in the catch by unit and 30% by weight.
Furthermore, these areas are nurseries for the migra-
tory species. Therefore, reduction in flooded forest
habitat would have broad impacts since many of these
species are found as adults in nutrient-poor black
water and clear water rivers. Loss of flooded forest
would reduce local fish production and that in nutrient-
poor rivers because of their dependence on recruit-
ment from the white water nurseries.
Our results suggest that common farming practices
on the floodplain may be in conflict with conserving
fish abundance. The abundance of fish is higher in the
flooded forest nearest to the river, yet these areas are
the first to be cleared to create space for buildings,
access ways, pastures, or gardens. Forest reserves,
when kept, are located mostly on the inland side of the
properties, where fish tend to be less abundant.
Acknowledgments This study was funded by the Spanish
CYTED program and additional funds from Brazil’s CNPq and
NASA. JERS-1 SAR imagery was provided through the Global
Rain Forest Mapping Project of NASDA, the National Space
Development Agency of Japan (now JAXA, the Japan
Aerospace Exploration Agency). Thanks to C. Granado-
Lorencio and A.Varella for comments on an earlier version of
the manuscript, to Michael Goulding for recent references and
comments about migratory species, to C. da Silva, Yossa, F. Chu
Koo, J. Sanchéz-Botero, M. Picanço, J. Penna, W. Dias, and
J. Marinho for their assistance during fish sampling and to the
crew of boats Veloz-I and Cap. Dario.
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