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Many fishes undergo seasonal migrations. In gen- to the seasonal flux of water level. Many fishes from
eral, migrations include longitudinal movements, floodplains exhibit morphological, physiological
along the main body of rivers, and lateral migra- and biochemical adaptations that permit their sur-
tions, between a river and its floodplain (Welcomme vival during periods of drought and anoxic water
1985). Many authors have reported longitudinal mi- (Junk et al. 1983; Saint-Paul 1984). Migratory be-
gration in characiform fishes of the seasonal flood- haviors may be a means to avoid such temporary ad-
plains of the Amazon Basin (Godoy 1979; Goulding verse conditions and they have been related to re-
1980; Worthman 1982; Ribeiro 1983; Zaniboni production and the utilization of allochthonous food
1985; Bayley & Petrere 1989; Menezes & Vazzoler resources (Goulding 1980).
1992). However, lateral migrations in the Amazon The objective of this study was to document and
have rarely been studied (Goulding 1980; Ribeiro discuss fish lateral migration patterns in a channel
1983). Daget (1958) and Welcomme (1985) charac- connecting a floodplain lake to the Amazon River. I
terized such movements for fishes of seasonally specifically address (1 ) the fish assemblage compo-
flooded African rivers. Here I present the first study sition of the study area: (2) patterns of lateral move-
of lateral migrations in the Amazon. ments of these fishes; and (3) detailed patterns of
Along the Amazon River, the seasonal water level migration of 6 species of Curamatidae and Prochilo-
fluctuation is quite large, with a mean variation of dontidae.
approximately 10 m per year (Junk 1983). Flood
pulse is driven primarily by seasonal precipitation Methods
cycles in the upstream catchment rather than by lo-
cal rainfall and is the major force regulating biota in Migration patterns and their physiological correlates
river-floodplain systems (Junk 1984; Junk, Bayley were assessed using (1) net captures; (2) direct ob-
& Sparks 1989). There is, however, little documen- servations of fish movements; (3) observations of
tation of specific movements of animals in response reproductive chronology; (4) catch records from
36
Lateral migration of fishes in Amazon floodplains
subsistence and commercial fisheries, and ( 5 ) asso-
ciations of migration behavior with water level. n I
30/2500
Study area "1 2000
5
3
This work was carried out on Careiro Island (Stern- 1500 ,E
berg 1956; Annibal 1983; Anonymous 1988), the -
VI
m
3
first island downstream from the confluence of the
Rios Solim6es and Negro (Fig. 1). The island is sit- lo00 2-u
uated at 3" 05' to 3" 12' S and 59" 35' to 59" 50' W -
E
37
Fernandes
can kill fish as they move through the channel (see The common cast net was fished alone for 1 h, or
also Annibal1983; Odinetz-Collart & Moreira 1989). both cast nets were used for 30 min each. Fish catch
Odinetz-Collart & Moreira (1989) characterized per unit of effort (CPUE) among nets was standard-
the seasonal limnology of Parana do Rei among ized to number of individuals per hour per 100 m2 of
other places in Careiro Island, in 1987. The maxi- gill net and to number of individuals per hour of cast
mum oxygen concentration is found in January, June nets.
and October (9-12.1 mg.l-'), while the minimum The direction in which fishes were migrating, in
occurs in March and at the end of August to the be-
ginning of September (1-3 mg - I-'). Daily variations
of the dissolved oxygen show that there is an in-
crease during the day, with the maximum between
2-8 pm, at the surface and at 2 m depth. Conductiv-
ity ranged between 46 pS-cm-'. 1-' in September and
79 yS-cm-'.I-' in December, and pH varied be-
tween 5.2 (July-August) and 7.1 (March). No signif-
icant seasonal variation was found in surface water
temperature (26.7-29.4"C).
Experimental procedure
Tagging methods, which are often used for the study
of migration in temperate fishes, have proven unsuc-
a s o n d j f m a m j j a s
cessful in studies of neotropical fishes (Godoy 1979;
Worthman 1982; Carvalho 1983). Petrere (1985)
suggested that the low recapture rates of Amazon
River fishes is related to high mortality and disper-
sion of fish across a broad area. In this study, migra-
tion patterns were assessed using net captures and
direct observations.
38
Lateral migration of fishes in Amazon floodplains
December (exiting or entering Lago do Rei), was @ Potornorhim bfiw @ Pwctropmtrr rwtiloidre
39
Fernandes
Table 1. Species present in the catch of the three gears in Parano do Rei. Numbers between parenthesis refer to the species family: (1) Osteoglossidae; (2)
Clupeidae; (3) Engraulidae; (4) Characidae; (5) Cynodontidae; (6) Hemiodontidae; (7) Erythrinidae; (8) Prochilodontidae; (9) Curimatidae; (10) Anostomidae; (11)
Apteronotidae; (12) Sternopygidae; (13) Rhamphichthyidae; (14) Doradidae; (15) Auchenipteridae; (16) Ageneiosidae; (17) Pimelodidae; (18) Hypophthalmidae;
(19) Cetopsidae; (20) Loricariidae; (21) Cichlidae; (22) Scianidae.
40
Lateral migration of fishes in Amazon floodplains
l? Latior; l? rutiloides, P. amazonica and C. kneri
Results
showed advanced stages of maturation (mature and
I captured 3,646 individuals of 87 species represent- prespawning) from November 1985 to April 1986
ing 22 families (Table 1). Of these, 20 species were (Fig. 4). In both September 1985 and 1986, none of
clearly laterally migrating. An additional 10 species these fishes exhibited ripe gonads, and several had
were probably also migrating, but they were not ob- spent gonads. S. taeniurus and I! nigricans appeared
served in sufficient numbers to be classified as mi- ready to reproduce in December 1985 and January
gratory. 1986. In September and October of 1985, most indi-
Migrations from the lake appeared to peak twice, viduals were spent or immature, although a few
as measured by CPUE (Fig. 3). The first movement were in maturing and prespawning stages.
occurred during the receding of the waters from Au-
gust to September 1985, when the currents in the Subsistence and commercial fishery data
parana were from the lake to the main river. The
second occurred from December 1985 to January The subsistence fishery yielded its highest capture
1986, when current of the water in the parana was between September and April of both years (Fig. 2).
from the Amazon to the lake, at the beginning of the Each year the catch was lower between May and
floods. Of the six focus species (Table 2), Potamor- August. Two peaks of capture occurred, with the
phina latior, Psectrogaster amazonica, Curimata pattern of abundance similar to the gill net and cast
kneri exhibited periods of increased abundance in net data. The highest capture rates were during the
August-September 1985 and December 1985, receding water in September 1985 and January
Semaprochilodus taeniurus and Prochilodus nigri- 1986. These data corresponded to my own captures
cans yielded low catches in August to September (compare Fig. 2 and Fig. 3).
1985 and peaks of capture in December 1985; and Fig. 5 illustrates commercial fishing capture in
Psectrogaster rutiloides showed the reverse with a the channel over a 10-year period, and also displays
high yield in August and total absence in December. the mean water level of the Rio Negro at Manaus. A
Between August 1985 and January 1986, fish comparison of fish catch and water level suggests
were observed schooling and migrating, from the that the catches are associated with the falling wa-
lake to the Amazon up-river. During December all ter/dry season period, and also with the migration of
individuals were captured on the lake side of the gill fish from the lake.
nets; for instance, during the second week of De-
cember, 249 individuals of the six focus species
Discussion
were captured leaving the lake, while no individuals
of these species were found to be entering the lake. This study suggests the following three-phase pat-
In January and February some dispersed non- tern of lateral migration in the channel of Lago do
schooling individuals of migratory species were ob- Rei.
served in gill net captures to be moving inward from ( 1 ) In August - September during the recession of
the river to the channel. waters, some groups of P. latior; 19 amazonica, l?
Table 2. Total capture per unit of effort (number of individuals per hour per 100 m2 of gill nets and number of individuals per hour of cast nets) for 6 focus species
using two fishing gears, between August 1985 and September 1986. Note that gill nets were not used in September 1985.
Months
Species Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep
Gill nets:
Potamorphina latior 588 - 0 0 743 22 3 7 0 0 0 0 0 0
Psectrogaster amazonica 0 - 0 0 36 3 0 3 0 3 0 0 0 0
Psectrogaster rutiloides 40 - 0 0 0 0 2 0 0 3 0 0 0 0
Curimata kneri 20 - 0 0 109 0 0 0 0 0 0 0 0 0
Prochilodus nigricans 0 - 0 0 42 0 0 0 0 0 0 0 0 0
Semaprochilodustaeniurus 0 - 0 0 174 21 0 0 0 0 0 0 0 0
Cast nets:
Potamorhina latior 6 32 0 2 25 25 14 9 6 0 0 0 0 8
Psectrogaster amazonica 1 10 0 0 1 1 0 1 4 0 0 0 0 1
Psectrogaster rutiloides 3 4 0 0 0 3 1 0 3 0 1 0 0 3
Curimata kneri 1 1 0 0 0 0 0 0 5 0 0 0 0 0
Prochilodus nigricans 0 3 4 0 1 0 0 0 0 0 0 0 0 8
Semaprochilodustaeniurus 1 1 0 0 0 0 0 0 0 0 0 0 0 2
41
Fernandes
were 2 m above the long-term average during the
low water and high water, these does not seemed to
have interfered with the migration behavior. Fish
seem to migrate every year, in different concentra-
tions, in spite of the intensity of the flood pulse (Fig.
5).
Many authors have observed migrations from
tributaries at the onset of receding water. Migration
behavior might be initiated by changing food re-
sources (Shulman 1974; Saldaiia & Venables 1983).
"EAR For example, some fishes have been shown to feed
Fig. 5. Biweekly seine net catch in the Parana do Rei, related for many months in the inundated forest before
with the level of the waters of the Rio Negro between 1976 and their migration (Goulding 1980; Goulding & Car-
1986. Water level from 1976 through 1986 based on daily valho 1982; Lowe-McConnell 1987). Maximum ca-
records for the Rio Negro, provided by the Administration of
the Port of Manaus (PORTOBRAS). Solid line refers to total
loric content of Prochilodus mariae in the Orinoco
fish catch; dashed line to water level. river occurs at the beginning of the migratory sea-
son (Saldaiia & Venables 1983), and in general
maximum fat content of whole fish may reach 16 to
30% during high water (Junk 1984). Young
rutiloides, C. kneri, F! nigricans, and S. taeniums Sernaprochilodus, abandoning floodplain lakes dur-
leave the Lago do Rei swimming in the direction of ing receding waters and entering the Rio Negro,
the Amazon River. The movement follows the same have been shown to be fattened from their feeding
direction of the currents of the channel. When the in floodplain lakes (Ribeiro 1983).
fish reach the river, they go up river against the cur- It is also possible that an oxygen gradient might
rent in the direction of the Rio Solimbes. also serve as a stimulus for fish to leave the lake.
(2) At the beginning of rising water (December - Odinetz-Collart & Moreira (1989) observed a drop
January), a second lateral migration occurs, appar- in the level of dissolved oxygen in the Lago do Rei,
ently related to reproduction, when the fish again during August - September 1987. I also measured
leave the floodplain lake and then migrate up the low levels of oxygen during the falling of water, in
river. These fishes are mature, and from this it is in- the channel and near the entrance of the lake. The
ferred that their migration is related to spawning. water of the Parana do Rei was low in oxygen, pre-
Since water in the channel is now flowing into the sumably because of the large quantities of organic
lake, this migration involves swimming against the material and detritus from the flooded terrestrial
current. vegetation available for decomposition. In addition,
(3) After spawning (January-May), a lateral mi- dense aquatic plants and the inundated forest had re-
gration entering the recently flooded plains occurs, duced the insulation in the surface of the water dur-
in a dispersed way, without formation of schools. ing flooding, also decreasing the production of oxy-
The currents of the river favor larval drift and swim- gen by the phytoplankton (Junk 1984). It is well
ming of spent adults through the floodplain which known that fish are capable of behavioral responses
presumably functions as feeding and nursery to the oxygen concentration of water (Smith 1985).
grounds. In 1986, I observed a large number of moribund in-
The first movement from the lake (August-Sep- dividuals of migratory species, moving from the
tember) does not appear to be related to reproduc- lake. Apparently these fish suffered from anoxia fol-
tion. The majority of individuals captured at this lowing an algae bloom, because they were breathing
time had non-mature or spent gonads, but some indi- in the superficial layer of the water. In general, sea-
viduals were in the initial stages of maturation, and sonal changes in depth, current speed, current direc-
all had accumulations of fat in the abdominal cavity. tion, turbulence, conductivity, dissolved salt and ox-
Once in the river these fishes initiate a longitudinal ygen concentration may also serve as migratory
migration up-river (Fernandes, personal observa- stimuli to fish.
tion). Goulding (1983) noted that the receding-water The lateral migration exhibited by part of the fish
period was the most productive for fishing on the assemblage leaving the system favors the survival of
floodplains. The same was observed in this work. the remaining component. Because of the decrease
The main families caught in the channel entrance at of the population density, the resources available to
that time were: Curimatidae, Characidae, Anostomi- the fish remaining will be greater. Some individuals
dae, Hemiodontidae and Pimelodidae (Table 1j. of these species might stay in the lake because they
Although the hydrological conditions in 1985-86 are not in a suitable physiological condition to mi-
42
Lateral migration of fishes in Amazon floodplains
grate (e.g., insufficient fat reserves). Their chances rios de seis especies de las familias Curimatidae y Prochilodon-
of survival in the lake during the dry period may be tidae: Potamorhina latior, Psectrogaster amazonica, I? ruti-
loides, Curimata kneri, Prochilodus nigricans y Semaprochilo-
improved because of the reduction of fish density. dus taeniurus.
Reproductive migration, e.g., the second pulse of 4. Parece que las migraciones laterales en este ecosistema esdn
migration here, have been observed at the beginning asociadas a la reproduccidn y 10s cambios fisicos en el hhbitat
of floods by many researchers (Ribeiro 1983; Junk que se producen como consecuencia de 10s cambios en el nivel
1984; Carvalho & Merona 1986). Fish that initially del agua.
remained in the lake, caught from December to
March, were found to exhibit reproductive maturity. Acknowledgements
This is consistent with the idea that when this sec-
I gratefully acknowledge the endless patience of Jeff Podos and
ond pulse of fish reach the river, their longitudinal John Lundberg in revising the manuscript, as well as input from
upstream migration is reproductive in nature. two anonymous reviewers. This work was part of a CNPq (Con-
It is likely that spent individuals migrate from the selho Nacional de Pesquisa e Tecnologia), INPA (Instituto Na-
river into the floodplain and the forests to rebuild cional de Pesquisas da AmazBnia) and ORSTON (Institut
their energy storage. As inferred from the low FranGais de Recherches pour le Development en Cooperation)
project, and was supported by the European Community (CEE).
CPUE, fish do not school as they re-enter the chan-
nel. Goulding (1980) and the fishers from the region
of the Rio Madeira also observed that spent fish do
not move in large schools, but move to the tributar- References
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