15.

The aquatic mammals and feptiles of the Amazon

R. C. Best

1. Introduction

The vast river system~, f100ded forests and floodplains of the Amazon have
provided the necessary ecological conditions fm the evolution of an interesting
and extremely diverse aquatic fauna. The magnitude of which is implied by the
2000 fish species (Roherts IlJT2 ), over lOO species of amphibians (Hödl, in prep.),
11 species of chelonian ami six species of aquatic mammals.
Pink fiver dolphins. giant snakes. caimans and manatees are weil known in
popular accounts uf the Amazon, yet our scientific understanding of the hiology
of any of these animals is rudimentar) . With few exceptions, the anecdotal and
descriptive accounts of the first explorers. missionaries and naturalists (Bates
1864: Verfssimo 18lJ5: Humholdt 1889: Goeldi 1893: Ferreira IlJ03: Wall ace 1853)
are the hasis of our current kllowkdgc.
Only recently has interest in such animals revived, yet there has been insuffi-
eient time to studie IJ1tcnsively am single species. In this chapter I will review the
hiology and present statu~ of the aquatic mammals and reptiles of the Amazoll
regIOn.

2. The mammaIs

2.1 Amazonian rnaflUrCC (Trichcchus inunguis)

The Amazonian Ill<lnatec is thc onl\ freshwater sireIllan endemic to the Amazon
Basin, where it ranges from the Ilha de Maraj() to the headwaters in Colombia,
Peru and Ecuador (Domning IlJ81. Thornhack & Jcnkins 1982). It has heen
suggested (Ronald cr af. 1978: Husar ILJ77) that this species mayaiso occur in the
Orinoco flVer drainage s\slem hased on a description by Humboldt & Weigmann
(1838). Howc\·cl. It Ilw; not ileen !ound In the upper Negro river (Best. un-
published) nor in thl' hcadwatcr, of tbc Orinoco river (Mondolfi 1974). lt is

Sioli, H. (cd,) Thl' AIll;JLO!l Lll11n()i()~\ ;lIHj ialld'il"lik' l'l'(ll()g~ (l! a !llighl~ trnplcal rivcr and its basin. ISBN YO-6193-10g-K.
© 19H4 Dr \\ junk Puhlishl'l"-. !)()rdr,-'cht, BOq(\ll L;llll"]'i [\.'r Pnntl'd itl thc ~dth,:rland;-;.
372

present in all the major tributaries of the Amazon drainage system, where its
distribution is apparently restricted by two factors: fast turbulent water (rapids)
and the lack of aquatic vegetation in such areas.
Manatees have long been astapIe food of the indigenous peoples and also of the
riverine caboclos of the region (Photo 1). As early as 1542 the manatee was

Photo 1 Trichechus inunguis (manatee, peixen boi), killed by a caboc/o. (Photo: Anonymous).
 373

recorded as astapIe dietary itcm (Medina 1934), and Acuiia (1859) and Bates
(1864) mention it as weil as river turtles as common foods, Commercial exploita-
tion of manatee meaL or mixira, as the finished product (produced by frying
pieces of meat in the blubber of the animal and storing it in thc fat) is called, was in
full production as early as 1580, anel in the mid-1600's Padre Vieira mentioned that
up to 20 Dutch ships filled with manatee meat were sent annually to Europe (Silva
1790). The Pescaria Real da Villa Franca, operating in the mid-1780's ne ar
Santarem , produced in two years 3873 arro/ws of dried and salted manatee me at
(1 arraba = 15 kg) and 1613 potes (pole = 20-30 kg) of lard, the equivalent of about
1500 manatees (Ferreira ]903; Domning 1982). In the period between 1935 and
1954 , a new era of commcrcial hunting came into practice as the tanneries
developed techniques allowing the preservation of the thick, strong manatee skin
which was used for machincry belting. hüsing. high-pressure gaskets and glue
(Mendes 1938; Pereira 1944). During this 20 year period, between 80000 and
140000 manatees were probably killed in addition to the subsistence kill in the
more remote areas of the Amazon (Domning 1982) (Photo 2). The hide industry
stopped , possibly as a result of thc increased use of synthetic rubber and artificial

Photo 2 Result 01' a single dav of manatcc hunting in Lake Aiapua . Rio Purtis. about 1940. (Photo:
Anonymous).
374

materials; but also by this time a serious reduction in the existing population had
probably occurred. Both commercial and subsistence meat hunting continued
until between 1967 and 1973 when this species was protected by Brazilian law as
an endangered species (it is also protected in Peru and Colombia but not yet in
Ecuador). Although there still exists small-scale illicit commercial hunting with
the meat being sold in the markets of Amazon towns, subsistence hunting
continues unabated in the more remote regions such as the J apura river, where 29
manatees were killed in a relatively small area in August and September, 1981
(Best, unpublished). The vast area of the Amazon drainage and the shortage of
game wardens (22 at present in the State of Amazonas) makes more effective
management impossible.
Biologically, manatees have been little studied, probably because of their
secretive nature and the difficulty of observation in the turbid waters of the
Amazon. In 1974, a study was initiated at the Instituto Nacional de Pesquisas da
Amazönia (INPA) with the objective of learning enough about the biology of the
manatee to be able to implement an effective conservation program to preserve
this interesting, endangered species.
Manatees have been placed in the superorder Paenungulata along with the
elephant and hyrax by systematists. Moreover biochemical evidence (amino-acid
sequences in the a-crystallin of the eye lens), (de Jong et al. 1981; de Jong &
Goodman 1982) as weil as alburnen systematics (Lowenstein et al. 1981) have
shown this to be correct and that instead of being derived from a common stern
group with the ungulates, they possibly represent one of the first offshoots of the
placental mammalian line. These unique aquatic herbivores may thus represent
one of the first mammalian invasions of the aquatic environment, they retain
'primitive' features such as an extremely low (36% 'normal') basal metabolic rate
(Gallivan & Best 1980). which is the principal factor in their excellent diving
capabilities (Kanwisher et al. in press), and a relatively labile core body tempera-
ture (Gallivan et al. 1983). Being entirely herbivorous, manatees eat a wide
variety of aquatic and semi-aquatic plants (Table 1) although grass es are probably
preferred foods (Best /981; Montgomery et al. 1981). They are non-ruminants and
have specialized hind-gut fermentation in a large paired caecum at the juncture of
the large and small intestine. Manatees consume about 8% of their body weight
daily with a digestive efficiency of between 45-70% depending on the fibre and
mineral content of the food plant (Best 1981: Best & Atkinson, in prep.).
The habits of wild Amazonian manatees in relation to the seasonally changing
environment are poorly known and Verissimo (1895) and Pereira (1944) have
summarized what is known by the fishermen of the region. With the recent
perfection of radio-tracking techniques (Best er al. 1981) it has become possible to
follow manatees constantly in their natural environment as an aid to field obser-
vations. Such studies of free-ranging Amazonian manatees are currently in
progress and preliminary results indicate that these animals are equally active
diurnally and nocturnally. When in feeding areas, they may move about 2.7 km/d
 375

Table 1 Aquatic plants eaten hv Amazonian manatees. Trichechus inunguis.

Aracea Poaceac (Gramincac)

Mantricardia arhorcsccnl Echinochlou palystachyu
Pistia slratioles Hl'rncnachne umplexicuulis
Convolvulaceae L.ecrsia hexandra
[parnaea aqualica L.lIziola spruceana
Operculina alala Orvza perennis
Fabaceae (Legummosae) Panicum purpurascens
Phasealus ovalus Paspalum fasciculalum
Lentibulareaccae P. repens
Utricularia f(,!iosu Pontederiaccae
U. rninor Eichhornia azurea
Moraceae E. crassipes
Cecropia spp. Polltederia cordata
Nymphaeaceae RCllssia ralllndifolia
Nyrnphaea sp. Salviniaceae
Si/lvillia allriculatu
S. minor
S. sprucei

Refercncc: Best 19k1.

(Montgomery et al. 1981). Behavioural studies using captive manatees show

similar results with the animals dividing thcir behaviour into: resting 17%,
feeding 33% and swimming 50% (Best. Ribeiro & Gallivan, in prep.). The
pronounced seasonal cycle of flooding in the Amazon drainage system has a
profound effect on the Iife cycle of manatees. As the water levels rise (December
to lune) there is a pronounced surge in aquatic plant production in the flooding
wirzeas and igapos (J unk 1970) and the manatees disperse into such areas feeding
on the lush new vegetation. During the dry season (July to November) the water
levels fall quickly. stranding most of the floating meadows leaving them inaccessi-
ble to the manatees. As the water levels commence to fall, the manatees initiate a
general migration trom the shallow floodplain areas to the deep water channels or
lakes where they pass the dry season protected trom the hazard of becoming
stranded. In 1963. one uf the driest years on record. hundreds of manatees were
killed by hunters as the deepest areas beeame shallow and imprisoned manatees
became vulnerable (Thornback & Jenkins 1982).
The lack uf obvious fond sources ami our observations of empty stomachs and
intestines of manatees killed at this time nf year suggests that the Amazonian
manatee may fast for an extended period 01' time during the dry season (Best
1983). They therefore must store sufficient energy during the wet season to be
able to maintain themselves during the next dry season. It may be that the highly
developed caecum is advantageous in maintaining such energy reserves, as it may
be important in deriving cnergy ami nutrients from ingested organic detritus on
376

the bottom of the lake through microbial fermentation in the caecum and colon
(Best 1981). The extremely low metabolic rate is also a critical adaptation for
survival during such prolonged fasting as it would extend considerably the time in
wh ich a manatee with good fat reserves could remain alive.
Manatees produce one calf per breeding after a gestation of approximately one
year. In the wild. the seasonal availability of food apparently restricts breeding to
the flood season when the females are able to attain sufficient energy stores to
sustain the energetic cost of pregnancy during the next dry season. Calving and
the more energetically costly lactation periods coincide with the period of highest
food availability allowing the female to regain her physiological condition (Best
1982). Newborn manatees are between 85-105 cm in length and weigh between 10
and 15 kg. In captivity they grow at an average rate of about 1-2 kg/week (Best
1982; Best er af. 1983).
The use of manatees to control aquatic plants is not a new idea (Allsopp 1960).
yet due to the difficulties of obtaining sufficient animals, little work has been done
to test their effectiveness (Nat. Acad. Sci. 1974). In 1980, a pilot study was
initiated at the hydroelectric dam of Curua-Una. ne ar Santarem (see Junk er af.
1981 for description of the area). To date we have introduced 42 manatees (20
females. 22 males) into the reservoir. each one radio-marked to be able to
determine the areas utilized by the manatees in relation 10 species and degree of
aquatic plant cover. daily and seasonal movements and also social behaviour, as
each individual is identifiable by the radio-frequency of its transmitter. It is
calculated that the 42 manatees in the lake will consume 20 metric tons/month and
a simple population model (Leslie matrix) has been made to project that in 11
years the population should attain 200 animals with a monthly consumption of
about 960 metric tons 01 plants.
The effect 01 manatees on the nutrient -cycles of lakes may be considerable
(Marlier 1967) if one considers that they may return to the lake over half the
material ingestcd in thc form of facccs ami urine. In the reservoir, that would
constitute about 330 kg daily of nutrients which are extremely rich (15% dTY
matter) in nitrogen (McLachlan 1971: Best. unpublished). Recent nutrient ad-
dition experiments in natural Amazonian waters indicate phytoplankton is nitro-
gen Iimited there (Zaret er uf. 1981) so that the excretory products of manatees
may be very important. Fittkau (1970. 1973) presents a similar hypothesis in
relation to the caimans of this region. However. the fact that manatees are
homeothermic. generally more active. and herbivorous suggests that they may
exert an even greater fertilization effect than the crocodilians. Certainly the
proposed increase in phytoplankton production resulting from the fertilization
effect of manatees should affect fish production in such an area (Melack 1976;
Lamarra 1975)
This work is continuing as a long term project with the aim of not only restoring
a more stable environment in hydroelectric reservoirs through the control of
aquatic plant growth but also as a mcans of initiating viable populations of the
 377

endangered Amazonian manatee in the large man-made lakes that are or will be
constructed in the Amazon region.

2.2 Giant otter (Pteronura brasiliensis) and the Amazonian river otter (Lutra
enudris)

Giant otters are the largest of the otter family, reaehing a totallength of 1.8 mand
weighing 32 kg (Photo 3). They are endemie to South Ameriea and have been
hunted almost to extinetion beeause of their extremely fine and valuable fur

Photo3 Pteronura brasiliensis (giant orter. ariranha) on shore with its fish prey. (Photo: Roberto H.
de Salvo Souza).
378

(Duplaix 1978; Smith 1981). In Brasil alone, more than 40 663 skins were exported
between 1960 and 1967 (An. Est. Brasil 1969). In 1972 they became a protected
species. Today they are principally rcstricted to thc remote areas of the Amazon
and Orinoco river drainages which have been inaccessible to hunters.
Although one of the most spectacular of the South-American mammals, this
species has been the subjcct of only a single ecological study, that of Duplaix
(1980). Additionally. there are so me short notes on their behaviour in captivity
(Autuori & Deutsch 1977; Salvo Souza & Best. in press; Trebbau 1972, 1978;
Zeller 1960). Virtually nothing is known of the river otter (L. enudris) (Duplaix
1978,1980; Harris 1968).
The giant otters inhabit small rivers, or streams (igarapes) which are usually
relatively dcar water. Groups of up to 20 individuals, have been recorded;
however. the normal group size is about 4-8 individuals, probably composed of
an adult pair and one or two current litters (Duplaix 1980). Such groups are
diurnal in their activity patterns and are extremely vocal, with a repertoire of
some 8-9 distinct souds (Duplaix 1980; Salvo Souza 1981). Their movements are
seasonaL varying with the changes in water levels of the rivers and creeks in which
they live as weil as the resulting availability of their prey (Duplaix 1980). In
Suriname, the trainl (Hoplias spp.) was eaten in over 50% of the observations,
with characoids and siluriform fishes being consumed less frequently. In our work
in thc Brazilian Amazon the arant (Schizodon sp.) is most common with Hoplias
spp. being less frequent (Salvo Souza & Best. unpublished).
The effect of both giant and river otters on the fish communities is probably
considerablc a~ mustelids tend to have high metabolic rates in comparison to
othcr mammals (I vcrson 1(72) and may consume up to 20% of their body weight
daily. Thus a group of six gwnt otters with an average body weight of 30 kg would
cat about 36 kg of fish dailv. an extremely high rate of eonsumption for the small
rivers in which thev live. Duplaix (1980) notes that such groups may travel
considerable distances and that thc size of their territories is related to food
abundancc.
Litter sizes of giant otters vary bctween one to six young, with a mean litter of
about 3 (Duplaix 1980; Autuori & Deutsch 1977; Trebbau 1972; Salvo Souza &
Best. in press). The gestation period is apparently about 70 days (Autuori &
Deutsch 1977;lrebbau 1(72). The young are born in river bank dens which are
used throughout the year. such dens being distributed throughout their range.
These dens ma~ often ha\c submerged entrances.
The much smalle! ri\CI otter is sympatric with Pteronura and may in fact
compete for f()()l1 rcsources at certain limes of the year; however. the greatly
disparate body sizes probahlv ensun:s that such competition is minimal. To my
knowlcdge, there IS not hing known about the biology of the Amazonian river
otter.
 379

2.3 Capybara (Hydrochoerus hydrochaeris)

Cabybaras are the largest living rodents (Photo 4) and range from Panama to
northern Argentina. These animals are amphibious and are seldom found far
from water. Although this species is of considerable economic value, both for its
meat and hide (Smith 1981: Ojasti & Padilla 1972), it has not yet been studied in
the Amazon region: however , in Venezuela it has been the subject of extensive
studies both of ecological and biological nature. Arecent study on social organi-
zation by Schaller & Cranshaw (1981) was made in the Brazilian Pantanal of the
State of Mato Grosso.
Capybaras weigh, on the average, about 49 kg and consume about 4 kg of fresh
grasses daily (Ojasti & Padilla 1972: Oiasti 1973 & 1983). Semi-aquatic grasses,
Hymenachne amplexicaulis, Leersia hexandra and Panicum spp. comprise the
bulk of their food intake although they do eat many other species (Table 2)
depending on season and availability (Ojasti 1973: Escobar & Gonzalez Jiminez
1976). The capybara digestive system consists oi" a simple stornach , along (~1O m)
small intestine with a large saccular caecum which contains about three-fourths of
the volume of the digestive tract (Ojasti 1983). The colon is about 3.5 m in length.
The digestive efficiency of capybaras is about 53'1'0 (range 50.9-64.5) for grasses

Photo 4 Capybara (J/vdrochoerus hvdrochaeris). thc largcst Iiving rodent (Photo: Anonymousl.
380

Table 2 Food plants of the capybara, Hydrochoerus hydrochaeris in Venezuela.

Gramineae (Poaceae) Pontederiaceae

Axonopus purpusii Eichhornia crassipes
Brachiaria plantaginea E. azurea
Cynodon daclylon E. diversifolia
Digitaria sanguinalis H eteranthera limosa
D. decumbens Marantaceae
Eragrostis pectinacea Thalia geniculata
E. hypnoides Boraginaceae
E. glomerata Cordia tetrandra (?)
Hymenachne amplexicaulis Papilionoideae (Leguminosae)
Leersia hexandra Machaerium aculeatum
Luziola pittieri Mimosoideae (Leguminosae)
Oplismenus burmannii Mimosa asperata
Panicum laxum Acacia glomerosa
P. zizanoides
Paratheria prostata
Paspalum orbiculatum
P. fasciculatum
P. plicatulum
P. repens
P. chaffanjonii
Reimarochloa acuta

Refercncc: Ojasti 1973.

which is similar to that of ruminants such as sheep (Ojasti 1973; Gonzalez Jimenez
& Escobar 1975). Stocking rates of capybara in open, savanna regions are about
1.8 animals/ha and in a forested habitat, 2.1 animals/ha (Cordeiro R. & Ojasti
1981; Ojasti 1980). Probably the value of2 animals/ha may be found in Amazonian
populations although this has not been investigated. The horne range of family
groups is about 10 ha (Ojasti 1980).
Although able to breed throughout the year, at a maximum rate of 2litters per
female, the capybara is reproductively synchronized to the periodicity of the
ecosystem. Most young are born late in the rainy season after a 5-month gestation
period. Females usually produce a single litter due to the seasonality of environ-
mental factors (Ojasti 1983). The peak in sexual activity occurs during the
beginning of the rainy season when water and new plants are abundant (Ojasti
1983). This type of breeding scasonality is identical to that found in the Amazo-
nian manatee (Best 1982). Average litter sizes are 4 young but up to 8 may be born
and they reach sexual maturity in about 18 months (Ojasti 1973; 1983). High infant
mortality (over 50%) is one of the major factors in regulating population levels,
especially during the dry season (Ojasti 1983). Natural predators sueh as the
jaguar (Panthera onca), caimans (Melanosuchus niger, Caiman crocodilus) and
anacondas (Eunecres mllrinus) have been less common in recent years due to
 3Rl

human predation and thus exert less influence on natural populations (Ojasti
1973, 1983; Schaller & Vasconcelos 197R). The great interest shown by South-
American countries such as Venezuela and to a lesser extent Brasil, Colombia
and Peru in the semi-domestication of the capybara (Bone 1977; Cordeiro 1977;
Cruz 1974; Fuerbunger 1974; Gonzalez limenez 1977; Zara 1973; Ojasti 1980;
Ojasti & Padilla 1972; Cordeiro & Ojasti 19RI; Macdonald 1981) has done much to
promote the study of the biology of this intercsting giant rodcnt.

2.4 River dolphins ((nia gcoffrensis und Sotalia fluviatilis)

The Amazonian freshwater dolphins represent two families of cetaceans. The

Amazon dolphin or bo!o vcrmelho (fnia geotlrensis) is a member of the primitive
family, Plastanistidae. which are typically freshwater in their distribution (Photo
5). Whereas the river dolphin or rucuxi (Soralia jluviafilis) is a member of the
Delphinidae , which are prcdominantly marine, fast swimming dolphins (Photos 6
and 7).
Inia is found in the Amazon and Orinoco drainage basins as weil as in the Beni
and Guapore rivers of Bolivia and there is so me controversy as to whether these
are different species. subspecies or. more simply, races (Pilleri & Gihr 1977 ;
Casinos & O<;ana 197<); Hershkovitz 1(6). A similar confusion exists for Sotalia

Photo 5 Ini" geo(f'rensi.\ (Amazon dolphin. holO vamelho) on land. (Photo: Vera M. F. da Silva).
382

Photo 6 Sotalia flllvialilis (river dolphin, llicuxi). (Photo: Vera M. F. da Silva).

for whieh five speeies have been deseribed (S. brasiliensis, S. fluviatilis, S. pallida.
S. guianensis and S tucuxi). Reeent authors (e.g. Mitchell 1975; Riee 1977)
recognize only S. fluviatilis. however the larger, coastal, Sotalia ranging from Rio
de Janeiro as far. north as Venezuela (Casinos et al. 1981) and Panama (Bössen-
ecker 1978) may in fact bc a different speeies. Riverine Sotalia are found in the
main tributaries of thc Amazon drainage as far as Ecuador and are also known
from Suriname (Husson 1978; Duplaix 1981) and Guyana (Williams 1928).
These two dolphins are very different in size. Inia reaching a length of 2.6 m
and a weight of 160 kg whiJc thc riverine Sotafia may be the smallest delphinid
attaining a maximum body length of 1.5 m and weighing 53 kg. Female Inia reach
,I
sexual maturity at bodv length of 183 em and produee a single ealf after a
gestation of aboLl112 months. The gestation period of the smaller Sotalia is about
10.2 months amI a 71 1075 cm calf is born. Female Sotalia are mature at body
lengtlls betwecn 121-\.'\ and 131-\.5 Clll (Best & Silva. in press). Both dolphins show a
breeding seasonality with the young being born as the water levels reeeed or when
 3i'\3

Photo 7 Sowliliflul'iatilis (lI/cl/xi), jumping. (Photo: Vera M. F. da Silva).

they are at their lowest levels, with Inia calving before Sotalia, Such seasonality is
a result of the synchrony of reproductive events with the time of greatest food
availability such that the energy cost of the last third of pregnancy, as weil as that
of lactation , may be offset by the ease of obtaining food energy, Although sam pIe
sizes are smalL it seems that the temporal difference in calving between the
species is related more specifically (0 food habits. The feeding ecology and
stomach content analyses of thc (WO species have shown that Inia feeds pre-
dominately on benthic. solitan . fishes while Soralia specializes in pelagic, school-
ing fishes (Silva 19R3) Thus. the fish that Inia preys upon would become vulncra-
ble as SOOI1 as the water leveL, start 10 receed and reduce the available cover in
igapo and I'arzea are(lS, whereds the pelaglC fishes preved upon by Sotalia would
be most vulnerable only as the waters reaeh their lowest levels (Best & Silva, in
press).
Another interesting aspect ol the reproductive biology of the freshwater dol-
phins are the contrasting male breeding systems. Inia males become mature at
lengths of over 199 cm and the testes weights increase proportionately with body
size. attaining a maximum size of 1.2'X, of body weight. Sotalia males, however,
re ach sexual maturity at a body length of 140 cm but have proportionately much
larger testes, up to 5'~J ofbody weight. Additi(mally, only so me of the adult males
have such greatly enlarged testicles. while the others have testes only slightly
larger than those of immature males (Best & Silva in press). In previous field
studies (Magnusson er al. 1980) we have shown that Inia is basically a solitary
dolphin and Sout/ia is more social. normally occurring in groups of 2-6 individu-
als. We interpret the very different testes ratios in the following manner: male
lnia probably resort to intraspecific combat to decide which male will mate with
an oestrus female. thus representing a monogamous mating system (Eisen berg
1981). Male Souilia probably form a breeding group around an oestrus female and
at the time of copulation the female accepts various males. In this case, it is not
the males that compete directly with each other but the sperm of each male is in
competition within the uterus of the female. It is thus to the advantage of the male
to produce more sperm than his competitor and hence the much greater testes size
(Best & Si/va. in press).
Inia and Sotalia are known to have the acoustic fat tissues in the forehead or
melon (Ackman el al. 1971.1975; Litchfield & Greenberg 1979) and are capable of
making communication as weil as echolocation sounds (Caldwell el af. 1966;
Caldwell & CaldwellI970a.1979b; Norris el al. 1972; Nakasai & Takemura 1975).
Echolocation was experimentally demonstrated in Inia in captivity (Penner &
Murchison 1970) and it is more than probable that both species use echolocation
as the primary means of prey detection in the turbid Amazon waters. Although
the related Indian river dolphin Platanista is blind, with degenerate eyes (Heraid
et al. 1969). Inia and Sotalia possess a reduced number of neurons in the optic
nerve. 19500 for Sotalia and 16500 for lnia. versus 150000 to 170000 in oceanic
species (Morgan & Jacobs 1972) but retains a relatively efficient visual ability
(Phillips and McCain 1964).
Silva (1983) is currently studying the food species eaten by these dolphins and
has, to date. found a total of 53 species of fish in the stomach contents. Of these
fish, 45 species were utilized by Inia and 28 species by Sotalia (Table 3). Fourteen
species of fish werc eaten by both dolphins with the freshwater croakers (Plagios-
äon spp.) being found in 57'10 of Inia stomachs and 33% of Sotalia stomachs. All
species of fish eaten by Sotalia were pelagic and some 75% were schooling fishes.
Only 58% of the fish eaten by lnia were pelagic in nature with about 42% of these
forming schools. A small river turle (Podocnemis sextuberculata) was found in the
stomach of one lnia (Silva & Best 1982) and in the Rio Guapore, Inia has been
reported feeding on crabs (Pilleri 1972). Large pectoral spines of doradid catfish
are common in lnia stomach contents and in most cases these large fish have been
tom into smaller pieces to allow ingestion (Silva 1983).
Most of our knowledge of the behaviour of the freshwater dolphins of the
Amazon comes from animals maintained in captivity (Spotte 1967; Huffman 1970;
 385

Table 3 Prey specics eaten by thc freshwater dolphins, lnia geoffrensis and So/alia fluvia/ilis.

Inia geoffrensis Pimelodidae

Cichlidae Brachyplalysloma filamenlosum
Geophagus surinamensi.\ Brachyplatys/oma flavicans
Gchla occelaris Pselldopla/ystoma /igrinus
Creniciehla lugubris Pinirampus pinirampu
Aearich/hys heckelii Pimelodus blockii
Astronotus ocellalus Sorubim lima
Sciaenidac Doradidae
Plagioscion spp. Megaiodoras irwini
Pachypops spp.
Engraulidae Sotalia Ouviatilis
L ycengraulis batesii Sciaenidae
Loricariidae Plagioscion spp.
Loricariichllzys acutus Pachypops spp.
Loricariinae Clupcidae
Hypophthalmidae Jlislza sp.
Hypoph/Izalmus perporosus Pellona jlavipinnis
Hypoplzllzalmus spp, Curimatidac
Sternopigidac ClIrima/a ciliala
Sternopiginae ('. laliar
Ctenolucidae ('. laliceps
Boulengerella spp. ('. viltala
Osteoglossidae Cilrimaleila meyere
Os/eoglossum bicirrhoswn Eigenmannifla meluflopogofl
Erythrinidae Prochilodus sp.
Hoplias malabaricus Anostomidac
Curimatidae Schizodon Iascia/us
Eigenmannina melal1opogol1 Leporinlls fascialus
Curimala la/ieeps Scrrasalmidae
C. la/ior Mylossoma aureum
Polamorhillu prisligasler M. dllrivellire
Hemiodidae Cichlidae
Hemiodus unimaculalll.1 Acarichlhys heckelii
Hemiadopsis microlepis Erythrinidac
Hemiodopsis immaClllallis f-{oplias malabaricus
Clupeidae Pi111c!odidae
Pellona cas/eillaeana ('al/ophvslIs macroplerlls
Serrasalmidac T'ime/odus hlackü
Colossoma bidens Pinirampus pinirampll
Serrasalmus (Pristobrvcoll) sp. A uchenipteridac
Mylossoma duriven/re Aucheniprerus sp.
Melhvnnis spp. Characidac
Anostomidac Rhaphiodofl villpinlls
Leporinus sp. H\'dro/ycus sr.
Laemoly/a varia Triporlheus spp,
Characidae Rn'('(m sp.
Brycon sp. Hypopthalmidae
Siluriformes Ifypophlhalmus perporosus
undetcrmined spccies

Reference: Silva 19iD

386

Caldwell & Caldwell 1969; Layne & Caldwe1l1964; Caldwell & Caldwe1l1972;
Caldwell et al. 1966; Gewalt 1979a, 1979b) and very few field observations have
been made of either species. Group sizes of Sotalia are generally between 2 or 3
individuals (55%) and rarely up to 9 individuals whereas, Inia is generally solitary
(81%) (Layne 1958; Magnusson et al. 1980). These data agree closely with those of
Pilleri (1969), Pilleri & Gihr (1977) for Inia in Bolivia and of Trebbau & van Bree
(1974) for Venezuelan Inia. In the Brazilian Amazon, on the Rio Solimäes,
Sotalia were the most common at a density of 1.6 individuals/km and Inia was to
be seen at a rate ofO.4 ind/km (Magnusson et al. 1980), while in Colombia, Layne
(1958) found lnia to be twice as common as Sotalia. This latter result more than
likely refects that lnia is able to utilize a larger variety of habitats, as it enters into
shallow varzea and igap6 areas whereas Sotalia remains in open water areas such
as river channels and lakes. Both dolphins are active day and night and they tend
to congregate at the mouths of rivers where the water is more turbulent. AI-
though Pilleri & Gihr (1977), Trebbau & van Bree (1974) suggest that Inia may be
territorial, this was not evident in our study (Magnusson et al. 1980). There is still
much to be learned of the ecology of lnia and Sotalia. Such studies are currently in
progress at INPA.

3. The reptiles

3.1 Side-necked turtles (Pelomedusidae and Chelidae)

The South-American river turtles of the genus Podocnemis are probably the best
known of the varied chelonian fauna of this continent. This group is considered
primitive and was one of the three reptile genera already in existence during the
Cretaceous period. 01' the SIX SOllth-American species, five are known from the
Amazon Basin: P expansa. P. wut/fis. P. erythrocephala, P. sextubercuiata and
P. dumeriliana. The iatter species has recently been accorded a separate genus,
Peltocephalus tracaxtl. due tn morphniogical. karyotypical, biochemieal and be-
havioural differences (Pntchard 1979a; Mittermeier 1978) as well as the fact that
[he hoiotype 01 P dumenliana is actuallv a P. unifilis (Pritchard 1979b).
Thc larger rivet' turtlcs. cspceiallv P expansa and P. unifilis, have been and still
are. principal foml ikms in lhe diet uf il1liigenolls peoples as weil as the mboclos.
As reviewed by Shrimpt!ln ()9f1()). and Smith (19 7 4.1979) the accollnts of ACllfia
(lfl59). Ferreira (19lUl. Ball" (181141. Coutinho OX6X). ami Sprllee (1908), early
exploitation of h\llh the egg\ dnd adult~ uf the various species of Podocnemis was
a highlv orgalllzed industn Srllllh 11979) documents the utilization of 208.9
million turtle el!g.\ fmrn historieal re ports on the Amazon. Ferreira (1903) men-
tions that in six ,car, I r;,~(~-17851, a total of 53468 turtles were captured by the
Portllguese Rll\C\J Fisherv 01 Barcellos (Rio Negro). Puraquequara (Rio Negro
near Manallsl dnd ManaeaPllni nr Caldenlo iRio Solim6es). The manner in
 3R7

which such turtle exploitation took place, included the protection of the tab-
uleiros or nesting beaches until the laying season had ended and then large parties
of people would gather the eggs, rendering them into butter which was used for
cooking and illuminating their houses (Bates 1864; Ferreira 1903). Very strict laws
were applied by the Portuguese to the use of the turtle beaches, and later
incorporated into the local municipal laws (Bates 1864; Pereira 1958; Ferrarini
1980). Adult turtles are still hunted throughout the Amazon, even though pro-
tected by the laws of 1967 and 1972, there is a ready market for them in the sm aller
interior towns as weil as in the city (Photo 8). Fishing techniques included baited
hooks (espinhel, camuri and cani{:o), harpoon or bow and arrow (jatica and
flecha) , traps (cacuri and paneiro) as weil as shooting them while sunning on logs
and catching the female as she is laying eggs (Ferrarini 1980; Smith 1979; Valle et
al. 1973; Ferreira 1903). The Brazilian Institute ofForestry Development (IBDF)
has been protecting some of the most important nesting beaches of the Amazon
since 1964, by placing wardens in the vicinity, thus preventing the capture of the
adult females and the ta king of eggs (Alfinito 1975 , 1978; Alfinito et al. 1973; Brito
1978; Alho et al. 1979). This protection has been expanded and includes nesting
beaches on the rivers Tapaj6s, Trombetas, Punis, Branco, Solimäes and Gua-
pore as weil as a number of privately preserved beaches.
Even though the degree of exploitation of podocnemid turtles has been consis-
tently high horn historical times to the present, extremely little is known of their
biology . Perhaps the most striking aspecL and certainly that which has received

Photo 8 Turtles confiscatcd froll1 a cOll1lllercial boat that was transporting Ihern 10 Manaus 10 be sold
illegally. (Photo: eicher .\Iho I
388

the most attention are the nesting habits and reproductive parameters which vary
considerably between species.
P. expansa is the only gregarious turtle during the nesting period and on
established nesting beaches or tabuleiros as they are locally called, as many as
6000 to 7000 turtles may use a single nesting beach during the three week laying
period (Alho & Padua 1982a; Vanzolini 1967). The nesting sequence of P.
expansa may be divided into about seven different phases of variable durations
(Alho & Padua 1982a, 1982b; Vanzolini 1967; Ojasti 1971; Padua & Alho 1982;
Valle et al. 1973). The first phase is that of the aggregation of the adults in the
deeper water ne ar the nesting beach. As the river levels receed, the adults are
seen more frequently in the shallow waters near the nesting beach, where large
numbers are visible with their heads oriented towards the beach. When the lowest
water levels have been reached, the turtles start to sun-bathe on the exposed sand
(Photo 9). They occasionally rub their heads in the sand or raise the head high as if
smelling the air. This behaviour may possibly be related to the sensing of the
environment al temperature (Alho & Padua 1982b) or the relative humidity of the
sand in relation to optimal conditions for oviposition. According to local fisher-
men, the sunning behaviour is a necessary prerequisite to the maturation of the
eggs in the females . The final stages of the reproductive sequence are the
exploration of the beach and the selection of the egg-Iaying site, the digging of the
nest, laying, covering the eggs (Photo 10) and returning to the water. The
synchrony of the laying cycle is intrinsically linked with the water levels of the

Photo 9 The turtles first spend same time sunbathing on the beach before laying their eggs. (Photo:
elcber Alho)
 3K9

Photo JO A femalc Podoel/emis eXpill1.l11 at the end of cgg laying . (Photo: elcber Alho).

rivers such that the tunIes in the Amazon nest between October and December
coincident with the low water levels (Alho & Padua 1982a; Vanzolini 1977). While
those in the Orinoco river do so in February or March apparently using the same
environmental cue (Ojasti 1971). Egg laying is usually nocturnal in the Amazo-
ni an and Orinocan P. expansa. yet the population on the Rio Tapaj6s nest during
the day (Valle el al. 1973).
A minimum carapace length (lf 50 cm is recorded for ovipositing females. and
the largest of the Tromoetas river P. expansa was 80 cm (Alho & Padua 1982a).
Ages are not knowll for these individuals and the only estimates of minimum age
of maturity are those of Ojasti (1971) who also cites Ramirez (1956), suggesting
that at least 5 to 7 years are necessary for this species to reach adult size. Alho et
al. (1979) have recorded individuals marked 15 years previously nesting on the
same beach. and the interval between nesting is about 4 years (Pritchard 1979a).
Hopefully the various long-term marking programs in progress in Brazil and
Venezuela will give more accurate estimates of such critical parameters. Re-
productive characteristics of the males are completely unknown although Alho et
390

al. (1979) have observed copulation in the shallow waters of the turtle beach
shortly after egg laying. The adult turtles remain near the nesting beach until
shortly after the young turtles have hatched (Alho et al. 1979).
The nest of P. expansa is hollowed out in the sand to a depth of 75-100 cm and
the egg chamber is about 13-18 cm deep and 20-25 cm wide. This excavation takes
a total of about 1-2 hours. the oviposition and covering ofthe nest taking about an
equallength of time (Vanzolini 1967; Alho et al. 1979; Ojasti 1971). There are
many reports on the numbers of egg in the clutch of P. expansa, these ranging
from 48 to 134 with a mean post ure of about 90 spherical parchment-like eggs,
each weighing trom 18-52 g (Alho & Padua 1982a; Ojasti 1971). Clutch size may
vary with the size or age of the females (Alho & Padua 1982a). The temperature
ofthe egg eh amber varies from about 29.5° C to 39° C (Alho & Padua 1982a; Ojasti
1971), the effect of such temperature variations on the sex determination of the
developing emhryos (Yntema 1969) has not yet been studied. Unseasonal flood-
ing (repiquete) may kill the developing embryos. The incubation period is short,
45-48 days with a hatching success of about 95°;;, under favorable conditions,
although unseasonal rising river levels may kill almost the total produetion if the
nests become t100ded (Alho & Padua 1982a; Mittermeier 1978). The young
turtles. about 55 mm in lcngth and 22 g. appear at the surfaee of the beach in two
waves. Firstly. about 60% of the hatchlings dig their way almost to the surface,
and three days later are joined by the rest and they leave the nest at night,
preferring rainy nights. There is relatively little predation of eggs in the nest,
although those eggs unearthed hy other turtles are eaten by vultures, terns and
teju lizards (Tupinambis nigripunctatus). Onee hatched the young turtles must
run a gauntlet of predators (Table 4) thay may eonsume up to 6'Yo of the hatehlings
(Roze 1974; Ojasti 1971). Certainly the colony nesting of P. expansa is related to
the survival value of synehronously produeing thousands of hatehlings during a
short period of time such that the predators cannot systematically eonsume more
than a small part of the overall production.
Knowledge 01' the biology of P. expansa while not on the nesting beach is
almost completely iacking. Some of the food plants have been deseribed for
Orinocan and Amazonian P. expansa (Tablc 5). The adult turtles are principally
herbivorous or frugivorous. yet it is probable that the young turtles eat propor-
tionally more protein-rich animal matter 10 aid their growth (Pereira 1958; Ojasti
1971) as recorded in other freshwater turtles (Clark & Gibbons 1969). The
stornach contents of 10 adult P. expansa killed in August and October contained
86% fruits and 4 (~,;) lcaves and stems (Ojasti 1971). The nature of turtle migrations
is unknown. However. it may be deduced from their food habits that they move
into the varzea and 19apo regIOns during the flooding season and as the waters
receed, the breeding adults must return to the nesting beaches (Ojasti 1971:
Pereira 1958). Non-breeding adults and juveniles probably remain in the larger
lakes (lago centraf) (Bates lR64: Pereira 1958) where there is often availab1e food
due to the landlocked nature of these lakes.
 391

Table 4 Predators 01' hatchling Amazon river turtks, Podoenemis spp,

Pisces
Piranhas Serrasalmlls spp
Catfish Hrll<'h\'plat\'sIOfllil, Phraclocepha!us, Rhamdia
Arawana (hl('og/OS,\'UII1 hicirr!Jo,\wlI
Pirarucu Am/Jilillla gigl/.\
Trair::i /loplll1s sI',
Peacock eich lid ( '{('il/a sp
Amphibia
Toad R/I[o sp
Reptiles
Caimans ('(///11(/11 cmcodilu,l,
,'vIc/"l1osuchus l1iger
Teju TllpilUllnbis nigripul1ClalU,\
Anaconda EUllenes lII11rIlIU,\

Aves
Raptors
Vultures Corag.l'p.\ alralll,l,
( alllllrfl'\' {lura
('ara-cara (arllCl/I'II /)11111111.\

Terns I'lwelll,\{/ simpl",

Herons and storks ./1I"il'/l IlIl'Clcria, Ardm herodias,
Luxenuril HWlf,uari
,\1r('/cria {/;}H'r/UlIlU

Mammalia
Amazon rivcr dolphin Ini({ geot}t·('lIsi.\
River otter 1,1/11'<1 sI'

References: Ojasti 1<)71: Si Iv<! 6: Best IlJ?-:2: Valle ('1111, 1973,

The other pelomedusid turtles have not been intensively studied and knowl-
edge of their biology is fragmentary, These species are solitary nesters with small
dutch sizes in comparison with p, expansa (Foote 1978; Neil1965; Mittermeicr
1978) (Table 6), Turtles of the genus Phrynops. by virtue of their very small
dutches (4-6 eggs). probably produce several dutches in different sites during the
course of several months (Medern 1960; Moll 1979), The remaining podocnemids
nest as the river levels receed and the bcachcs become prominent. p, tracaxa lives
in the sm aller streams and igap6 and nests in the vegetation along the banks as do
the species of Phrynops that inhabit the same environment (Medern 1960; Mitter-
meier el al. 1978; Pritchard 1979a)
The food habits of these turtles are gene rally unknown, Belkin & Gans (1968)
report on an unusual feeding mechanism of p, unifilis in which this species is able
to feed on particulate matter tloating on the surface of the water. P. expansa was
unable to feed in this manneL In captivity. p, unifilis readily feeds on aquatic
plants such as Eichhorniu a~ weil as fruits and meat, apparently being quite
omnivorous, The smaller P ervthrocephu/u and P. sextuburculata are mostly
392

Table 5 Food habits of the giant Amazon river turtle, Podocnemis expansa.

Leguminosae Moraceae
Campsiandra comosa Brosimium sp.
Macrolobuim acasieafolium Naucleopsis caloneura
Inga spp. Dillenracea
Tachigalia paniculaw Curutella americana
Palmac Simarubaceae
Bactris sp. Simaba guianensis
Astrocarym murWnllrtl Bignonaceae
A. tucllma Couralia toxophora
A. janari Rubiaceae
Gllilielma specioSl/ Genipa americana
DeSmOIJCIlS sp. Duckedendraceae
Compositae Duckeodendron cestoides
Mikania congesta Euphorbiaceae
Lecythidaceae Hevea brasiliensis
Jllgastrum sifolllesi H. spruceana
Gustavia augusllI Anacardiaceae
Cariniana sp. Spondias lutea
Sapotacea Verbenaceae
Lucuma lasiocarpa Vitex ounocensis
Micropholis sp. Elaeocarpaceae
Guttiferaceac Conceveiba guyanensis
Symphonia globulifera Humiriaceae
Callophyllllm sp. Saccoglottis guyanensis

References: Alho el ul. 11)79; Ojasti 1971: Pereira 1958.

herbivorous, feeding on plants and fruits of the igap6 and varzea, respectively
(Mittermeier & Wilson 1974). P. rufipes appears to be essentially herbivorous
although they eat small shrimps and fish, as weIl as palm fruits in the wild (Friar
1982; Medern 1973), and fish or snails in captivity. The remaining species, P.
(Mesolemmys) gibbus, P. (Batrachemys) nasutus and P. tracaxa are all predomi-
nantly carnivorous, easily being attracted to baited hooks, or by the smell of fish
chopped up in the water. A large captive P. nasutus would capture and eat live
white mice with an efficiency that rivalled the caimans in the same tank (Best,
pers.obs.).
The remaining ehe lids of the Amazon are both quite distinct for very different
reasons. Platemys platycephala is one of the smallest of the Arnazonian turtles,
and is the only chelonian of the region with a yellow or orange head. These turtles
are essentially aquatie, but often leave the water during rainstorrns to wander
sometimes a fair distanee frorn the nearest forest strearn or lake. They are
prirnarily earnivorous in eaptivity, feeding on meat, fish and snails. As they are
very agile in the water they are probably eapable of eatehing shrimps, sm all fish as
weIl as tadpoles. They nest all year long, produeing 4-6 eggs per cluteh, often
so rne distanee frorn water.
 393

Table 6 Some reproductivc parameters of Amazonian turtles.

Colonial Clutch Nestings

solitary size per year

Podocnemis erythrocephala S 5-14

P. expansa C 48-134 <1
P. sextuberculata S 6-16 1
P. uni/ilis S 21-40
Peltocephalus tracaxa S 7-25
Phrynops (Bactremys) nasuta S 2-4 >1
P. (Mesoclemysl gibbus S 2-4 >1
P. ru/ipes S ?
Platemys platycephala S 4-7 >1
Chelus /imbriatus S 12-28 1
Kinosternon scorpioides S 2-4 >I
Rhinoclemys punctularia S 2-4 >1

References: Mittermeier 1978; Mittermeier et al. 1978; Pritchard 1978a; Alho & Padua 1982. See also,
Medern. F. 1983. A reproduccion de 1a tortuga 'cabezon' Peltocephalus tracaxa (Spix), 1824. (Test-
udines, Pe1omedusidae). en Co1ombia. Loziana (Acta Zoo!. Colombia) 41:1-12.

The most distinct of the Amazon chelids in the mata-mata (Chelus fimbriatus)
with its prominent keeled shell, and strangely shaped head and neck. The mouth
is extremely wide and when opened rapidly it sucks in any unwary fish in the near
vicinity (Pritchard 1979a). The wart-like protuberances, are actually sensitive to
slight water movements such as those made by fish, and probably aid in the
detection of prey in the dark waters of the igap6 or forest streams (Hartline 1967).
Nesting in the Amazon is in October, when 12 to 28 round eggs are laid near the
edge of the stream (Medern 1960). Incubation time is approximately 208 days
(Hausmann 1968).

3.2 Musk turt/es (Kinosternidae) andfreshwater turtles (Emydidae)

There are only two species of aquatic turtles in the Amazon basin that retract the
neck directly into the shell instead of turning it sideways. The mussuä (Kinoster-
non scorpioides) is commonly sold house-to-house in Belem where it is a popular
dish even though illegal. The carapace of this species is elongated and domed with
three low kccls running longitudinally along the dorsum. The most distinctive
character are the hinged plates on the plastron. which dose against the carapace
completely protecting the soft parts against predation. The head is propor-
tionately large for this smalI. 10-15 cm turtle and they apparently feed on animal
matter (snails and insects) as weil as plants (Pritchard 1979a). In captivity they
feed on raw fish. They lay a single. elongate. hard-shelled egg. with the egg being
semi-buried in the lcaf-litter. Incubation takes between 126 and 148 days (Prit-
394

chard 1979a). Nothing is known as to breeding season although copulation has

been observed where the male holds the sheIl with all four feet as weIl as the
hooked tail (Sexton 1960). A subspecies, K.s. carajasensis has been described
from the Serra dos Carajas in the State of Para (Cunha 1970), although Pritchard
(1979a) regards this and other subspecies as being weakly characterized.
Rhinoclemys punctularia is found only in the lower portion of the Amazon
(Amapa and Para, as far west as Santarem and Parintins). Virtually nothing is
known of its biology although it appears to be semi-terrestrial. In captivity they
appear to be gregarious and they prefer shaIlow water. They feed on raw fish,
meat and aquatic plants, often fighting for morseis. This genus is notable for its
extremely large oblong eggs (-7 cm x 4.0 cm (Ewert 1978) laid in a clutch of
about 3, in a shaIlow depression covered by leaves (Pritchard 1979a; Medern
1962). Little more is known about this attractive turtle.

3.3 Snakes

Of the many species of snakes known from the Amazon region (53 species in the
Ecuadorian Amazon (DueIlman 1978) and 79 species at Belem (Cunha & Nasci-
mento 1978), only a smaIl proportion may be considered aquatic or semi-aquatic.
Apart from the weIl known anaconda (Eunectes murinus) , the largest snake of the
region (Amaral 1949), snakes of the genera Helicops, Hydrops, Hydrodynastes
and Micrurus are the most typical of this habitat type. These lesser snakes have
been little studied and their aquatic habits are mostly known from coIlection
localities. The known food habits are presented in Table 7. Helicops spp. may
either be viviparous or oviviparous. producing between 7 and 12 embryos or eggs

Tahle 7 Food habit, ()f thc water '!lake" lie/icops. liw/rops. liydrodynllsres ami Micrurus in the
AmaZOIl region.

Species I'rn

Helicops IIl1gU/lIll1,\
H. perersi fadpoks. small fish. lizards
H. po/r/e,,!.\ \ .\clIsricurtl.\) anel frog'

I fydrop.1 marlii Frcshwatl'1' ccl \Svmhranchus sp.)

!I. Iriangu/llri.1

Hvdrodvna.11Cs hicin(fll.\

,ViCrllrU'l ,,{{rilllllnCf!\j\ t::Jcctric ccl (G\'IIlIloles ('ampo).

,1fI11ourcd catfish (Clllliclhys
,'o/liel/n,l) and frcshwater cel
\Srlllhmnclills marmorallls)

Rcfercnccs: Dixon &: SOlni 1')77' ('unha & Nascimento [lJ7R: Duellman [lJ7S.
 395

with fully formed embryos ready to hatch (Cunha & Nascimento 1981; Duellman
1978).
The enormous anaconda, or sucuri as it is locally known, is the largest and most
feared of the snakes of Amazönia (Photo 11). There are many stories about its
attacks on humans and livestock and the mythical cobra-grande, or giant snake of
the Amazon, is in reality the anaconda suitably embellished for storytelling. The
actuallength of the largest Eunectes is a disputed fact, as certain authors mention
individuals of 20-22 m; however, scientifically verified records give maximum
lengths of 11.4 to 12 m (Pope 1961: Amara11949; Santos 1955). To my knowledge
there are no studies of the biology of this species and there is scant data on food
habits and reproduction. Anacondas feed principally on aquatic or semi-aquatic
animals such as fish, turtles , caimans, capivaras, deer, tapirs and aquatic birds
(Santos 1955 ; Haverschmidt 1970; Cunha & Nascimento 1978; Bellnomini et al.
1976177; Medern 1981 ; Ihering 1911). Most hunting is done nocturnally and the
prey is killed by constriction and preferentially ingested in the water. Feeding is
sporadic, as in most carnivores, and a captive female anaconda (3 m; 33 kg) ate 24
times in a 21 month period , consuming a total of 47.5 kg of rats, guinea pigs,
rabbits, pigeons, chickens and geese (DeschaneI1978). This amounts to about 2.3
gm/kg/day for an adult anaconda, whereas young growing anacondas require
about 6.1 gm/kg/day (DeschaneI1978). Mating takes place in the water , and the
gestation is about 270 days. Litters of anacondas vary from 14 to 82 live-born
young which are normally about 75 cm in length and weigh 200 gm (BelInomini et
al. 1976177; DeschaneI1978). Within a year the young have doubled their length
and in 3 years have attained a length of almost 3 m after which they grow more
slowly , but become proportionally much heavier (BelInomini et al. 1976177).

Photo 11 Big Eunecles murinus (anaconda . sucurf o r sucuri;ti) on the water surface in Rio Acre .
(From a postcard bought al Manaus ahout 1<J40).
396

3.4 Lizards and caimans

This last group of reptiles includes several genera of aquatic or semi-aquatic

lizards as weIl as four species of crocodilians, or more specifically, caimans. As in
previous accounts the existing literature on habits and ecology are extremely
scanty even for commercially exploited species such as the black caiman (Melano-
suchus niger). The taxonomy of most of these species has been studied in detail
(Cunha 1961; Hoogmoed 1973; Dixon & Soini 1975; Duellman 1978) yet only a few
studies have looked at the ecological relations and in these studies, the aquatic
lizards are usually poorly cited or absent (Rand & Humphrey 1968; Crump 1971).
An interesting fact is that the three most aquatic of the lizards (Dracaena,
Crocodilurus and Neusticurus) all have morphological characters that are remi-
niscent of the crocodilians, such as the keeled scales on the body of Dracaena and
the scutellation of their tails forming dorso-Iateral keels merging into a single keel
as the tail becomes narrow (Vanzolini 1965; Cunha 1961). This is probably a case
of convergent evolution in adapting to the aquatic environment and would be an
interesting subject to pursue further.
The most striking of the aquatic lizards is the large Dracaena guianensis or
jacuruxi as it is locally called. This large lizard reaches a length of over a meter and
has a distinctively scaled body that is, at first glance, easily mistaken for Caiman
crocodilus. Even though it has strong j aws with large blunt teeth (Vanzolini 1965)
it is a predator of aquatic snails, apparently eating few other prey items (Rand
1964; Vanzolini 1961; Conant 1955). Foraging is done submerged while the lizards
walk slowly along the bottom of the pool, po king their head into the fallen leaves
along the bottom, extending the tongue frequently. When a snail is found it is
brought to the surface and the shell is crushed with the strong blunt teeth, the
hard fragments being expelled by the tongue, the snail's operculum is often
ingested (Vanzolini 1961: Rand 1964). There is little more information about this
species. except that it swims weil and when not active, foraging, it may spend
large amounts of time. perched in trees over the water. Goeldi (1898; 1902)
apparently found a nest lodged in a termite nest and this is the only known
statement on the reproductive biology of this species, and Dixon & Soini (1957)
re port a female that had 17 yolked ovarian follicles. Interestingly, this lizard was
hunted extensively for its meat and skin, the latter being used in the making of
shoes, purses. belts and other leather goods (Cunha 1961; Carvalho 1967).
Very little is known about the biology of either Crocodilurus lacertinus, the
jacare-rana or Neusticurus ecpleopus or N. bicarinatus (UzzeI1966; Cunha 1961;
Duellman 1978; Goeldi 1902). Both species are apparently diurnal and swim and
dive easily. Mature females of N. ecpleopus range from 53 to 58 mm snout-
to-vent-Iength (SVL) while the males are slightly larger (52-64 mm SVL) (Dixon
& Soini 1975). Totallengths of Crocodilurus are over 50 cm while N. ecpleopus
reaches a totallength of about 17 cm (Duellman 1978; Goeldi 1902). Neusticurus is
restricted to small forested streams where it feeds on orthopterans, spiders, ants,
 397

beetle larvae, centipedes and other lizards (Anolis, Leposoma and Neusticurus)
(Duellman 1978). A single or two leathery eggs are produced and breeding is
apparently unseasonal in N. ecpleopus and N. bicarinata (Duellman, 1978;
Uzzell, 1966).

3.5 Ca imans (Melanosuchus. Caiman and Paleosuchus)

The caimans of the Amazon were once extremely common (Bates 1864; Pereira
1944; Fittkau 1973; Hagmann 1909; Wallace 1853). However, in the 1940's the
leather industry initiated a heavy exploitation of the black caiman (Melanosuchus
niger). Prior to this the caimans were occasionally shot or killed as pests around
houses or near fishing camps (Bates 1864). In Tefe. a small town on the Rio
Solimöes, oil rendered from caimans was used with diesel oil (70% and 30%,
respectively) to run the generators for lighting. Caiman oil was worth CR$15 per
tin and diesel was CR$55 for the same quantity (Pereira 1944). By the late 1950's
Melanosuchus was becoming extremely rare in all its range (Medern 1981; Smith
1981) and the tanne ries had learned how to tan the skin of the smaller spectacled
caiman (Caiman crocodilus) which was then hunted along with Melanosuchus.
Although prohibitcd in 1967 in Brazil and in 1968/69 in Colombia, the clandestine
hunt continues even today (Medem 1981; Smith 1981; Rabelo & Magnusson 1983).
The magnitude of this hunt may be gauged by the records far Caiman in Colombia
which amount to 10 million hides between 1951 and 1976 (Medern 1982). Probably
the kill of Melanosuchus was proportionally much greater as this species was the
most common in pre-exploitation times (Bates 1864). Both Melanosuchus and
Caiman are considered to be 'endangered' and 'vulnerable', respectively, by the
IUCN (Groombridge 19K2). A rccent devclopment in the killing of caimans is
that of the dried and salted me at trade in which all species are utilized including,
for the first time, thc smaller caimans of the genus Paleosuchus. The meat is
exported illegally. along with similarly prepared catfish meat, either to Belem or
Leticia (pers. obs.). For further data on the trade in Caiman skins, the reader is
referred to IUCN (1982b), IliCN (1973) and Medem (1981). Although caimans
have played an important part in the economy of the Amazon during the past 35
years. virtually 110thing has been done with regards to the study of their biology.
The black caiman or jacare-a~'zi (M. niger) attains an adult length of 6 m;
however. due to the extreme hunting pressures, such large individuals are ex-
tremely rare. This spccics prcfcrs quiet watcrs such as varzea lakes and igap6
forest, showing little preference far riversides where the water eurrents are faster
(Medern 1981: Magnusson 1980). This species was once extremely common in the
grassland (varzea) lakes of Ilha de Maraj6 (Hagmann 1909) and could be seen
along the sand bars of the rivers probably during the dry season when the
secondary lakes became too dry to sustain them. Juvenile M. niger feed on small
fish, amphibians, insects. crustaceans and snails. The large size of this species
39R

Table 8 The food habits of the hlack caiman, Melanosuchus niger.

Mollusca Reptilia
Gastropods Podocnemis unifitis
Bivalves P. expansa
Insecta Caiman crocodilus
Coleoptcra M. niger
Hemiptera Corallus caninus
Hymenoptera Lachesis muta
Odonata BOlhrops alrox
Orthoptera various Iizards
Isoptera Avcs
Arachnida Neoehen juhala
Spiders Cairina moschata
Crustacea Ammides cajanea
Crahs Lalerallus melanophais
Shrimps Porphyrula marlinica
Pisces Podiceps dominicus
Pimelodus sp. Phalcrocorax olivaceL/s
Cie/aL/rus sp. Casmerodius albus
Plecoslomus sp. ]acana jacana
Plagioseioll sp. Helicornis fulica
Prochilodus sp. Mammalia
CvnopOlamus sp. Didelphis sp.
Brvcoll sp. Chironecles sp.
Cvnodofl sp. Hvdrochoeris hydrochaerus
Serrasalmus sp. Dusyprocla sp.
Amphihia C/lniculus pa ca
Buto marinus Mazama, sp.
B. gral/ulosa
Eleulherodaclyltn ockendeni
E. ,·enlrimarmOI'llIll.1
Eda/orhil/a per<':!
LeplOdactv/us ho!i"iallus
Bo/ilhog/o.\.I'a allamazoilica

Rcfcrcnccs: Ottc !LJ7K: Medern ]LJi\1.

enables it to eat a wide variety of animals (Table 8) and, in fact, in areas such as
I1ha de Maraj6 they were even kilJed, as pests as they preyed upon young cattle on
the ranches (Santos 1961), Melanosuchus appears to be relatively gregarious, as
they were often seen lying together sunning (Bates 1864). In the lower Amazon
these large caimans may estivate during the dry season (Bates 1864; Hagmann
1909),
The reproduction of M, niger is poorly understood, with most of our present
knowledge coming from the works of Hagmann (1902; 1909) based on observa-
tions made at the mouth of the Amazon , The nest is a mound of leafy material of
about 80 cm in height and 150 cm in width, the eggs being buried at a depth of 40
 399

cm. The eggs are deposited in two layers with a layer of rotting vegetation in
between the layers. Average dutch size is from 30-60 hard shelled eggs that are
86-96 mm in length and 52-56 mm in width (Hagmann 1902; Goeldi 1898; Donoso
Barros 1966). Incubation is between 5 and 6 weeks, being less in nests which are
exposed to direct sunlight and more in those in shaded localities. The females
guard the nests, while incubation is in progress and respond to the distress calls of
the newly hatched young, a fact that has been exploited by skin hunters who easily
imitate this sound calling the female doser and then killing her. Breeding in the
lower Amazon coincides with the low water period of October and November.
Females of over 2 m body-length may be reproductively mature (Brazaitis 1974).
Captive young M. niger may grow between 35 and 50 cm/year (Donoso Barros
1966; Dowling & Brazaitis 1966). Further information on this species may be
found in Otte (1978) and Plotkin et al. (in press).
The spectaded caiman (e. crocodilus) is still relatively common in the
Amazon region in spite of the tremendous hunting pressure from the skin trade.
This very adaptable species was not hunted until about 1950 as the skins were
considered of poor quality by the tanning industry (Medern 1981). lt was pre-
viously sought after in the State of Para for its meat which was often sold as dried
pirarucu (Arapaima gigas) meat there. M. niger meat was not considered good to
eat because of its strong smell (Pereira 1944).
e. crocodilus are smaller than M. niger attaining a maximum body length of 250
cm and average lengtlls of 150-220 cm, and weigh up to 62 kg (Brazaitis 1973;
Staton & Dixon 1975). They are sexually dimorphic with the male being larger.
Spectaded caimans are found in many types of habitats, particularly along river
channels and lakes (Magnusson 1980; Medern 1971; 1981; Dixon & Soini 1977;
Vanzolini & Gomes 1979; Staton & Dixon 1975). They do not live in dosed forest
streams, yet may use them as routes through which they colonize inland lakes
wh ich have small outlets (Magnusson 1980). Like M. niger, this species is gregari-
ous, Iying dose together often touching while basking in the sun. Historically, e.
crocodilus was much less common than M. niger (Bates 1864; Hagmann 1910) and
probably the relative abundance was one C. crocodilus for 20 M. niger (Fittkau
1973). Today, after extensive commercial hunting, this ratio has been reversed
and in a sampIe of 1093 confiscated skins, there were 9 Caiman for each Melano-
suchus (Rebelo & Magnusson 1983).
Feeding studies by Fittkau (1973) have revealed that this species eats about 0.6-
0.8% of the bodyweight per day, which would be a daily intake of 0.4 kg/d for a
large 62 kg Caiman. Logically, bv being a predator, the feeding bouts would be
sporadic and the amount of food consumed in a given feeding would be much
greater. The food habits of young e. crocodilus differ from those of juveniles or
adults and are principally aquatic insects (Staton ]976; Staton & Dixon 1975).
Older specimens feed on a wide variety of foods (Table 9) normally ingested
whole. The peculiar occurrence of gastroliths (usually sm all stones) is common
(32.5% frequency) in this species (Staton & Dixon 1975). Caiman is also a carrion
40()

Tablc 9 The food habits of thc spectacled caiman, Cuimall crocodilus.

Insecta Anndidac
Odonata Worms
Orthoptera Mollusca
Hcmptcra Gastropoda
Coleoptera Pomacea
Diptcra Amphibia
Lepidoptera Hylidae
Hydrophilidac Lcptodactylus sp.
Dytiscidae Pisces
Belastomatidac Cichla.loma sp.
Arachnida Symbranchus sp.
Spiders Hoplia.l malabaricus
Platyhclminthes Aves
Lccchcs Todirostrum sp.
Crustacea Mammalia
Crah C' Tricl!oductvlll.l PCruvillllll.l) Rodentia

References: Staton &: Dixon 197'i: Vanzolini &: Gomcs 1979.

eater and cannibalisrn is a cornrnon phenornena related to population density

(Staton & Dixon 1975; Medern 1981). Interestingly, Caiman is able to feed on the
poisonous skinned South Arnerican toads, Bufo marinus and B. granulosa (Gor-
zula 1978; Medern 1981).
Physiological studies have shown that Caiman adjusts its behaviour in relation
to the arnbient temperature. When water temperatures are high (29-30° C) they
spend long periods subrnersed in it. If the water temperatures are lower, they take
to basking in sunlight during the daytirne and rnaintain their core temperatures
around 30-33°C whilst at night time it drops to 26-30°C (Groornbridge 1982).
In cornparison to the detailed work on reproduction done in Venezuela on this
species (Staton & Dixon 1977; Staton 1976), !ittle has been done in the Amazon
region (Hagmann 1906; Medern 1960; 1966; Chirivi-Gallego 1973). Reproduc-
tively adult females are about 1..2-1.3 rn in length at an age of about 4 years
(Chirivi-Gallego 1973; Staton & Dixon 1977). Copulation occurs in thc water.
Nesting is apparently seasonal and in Venezuela, coincides with the period
shortly after the rainy season (Staton & Dixon 1977), and at the rnouth of the
Amazon, Hagmann (1906) reported it to be in May and lune which is at the peak
of thc flood scason. In Colornbia (Rio Caguan) a nest was found in rnid-March
(Medern 1969). The nesting period is thus relatively flexible and allows the young
to hatch as the water levels of river or lake are high (and less !ikely to flood the
nest) and there is adequate food and cover for the hatchlings (Staton & Dixon
1977). Caiman nests are built at night using the available vegetation such as
grasses, twigs, leaves and earth. The eggs are deposited in the egg charnber which
is hollowed out after the ~ I rn x 1 rn x 20 crn nest pile has been made (Staton &
 401

Dixon 1977). The nests are placed near the water (20 m; Medern 1966) and the
female actively defends the nest against predators such as the teju lizard (Tupi-
nambis tequixin). Clutch sizes are variable (17-38; x= 29) and possibly varies
with female size (Staton & Dixon 1977; Chirivi-Gallego 1973; Medern 1958). Eggs
are elliptical (64.8 mm x 40.7 mm) weighing 59.9 gm and are hard-shelled.
Incubation is about 73 days at average nest temperatures of 28-32° C (Staton &
Dixon 1977). Nest predation accounts for between 50 and 84% of the prenatal
mortality (Staton 1976; Staton & Dixon 1977). Vocalizations of the young in the
nest stimulated the guar ding female to aid the young in removing them from the
eggs and taking away the eggshells.
The young are between 9.8-12.3 cm and weigh an average of 41.5 ± 0.6 g.
Predation on the newly hatched young accounts for about 83% mortality and
under crowded conditions cannibalism may elevate the mortality rates even more
(Staton & Dixon 1975; Staton 1976). Predators of this species are listed in Table
10. Growth rates of Caiman are about 30-35 cm/year (Chirivi-Gallego 1973).
The ecological effects of the great reduction of Melanosuchus and Caiman
populations are poorly understood due to the complex nature of the ecosystem.
Fittkau (1970, 1973) studied the effect of the large caimans in relation to nutrient
recycling and overall production in Amazon lakes. He extrapolated that caimans
released about 0.2 or 0.3°/" of their bodyweight daily as available nutrients (N, P,
Ca, Mg, Na and K) as excretory byproducts. As much of the caiman food is of
allochthonous origin. at pristine population levels (estimated to be 4 x 104 kg/km 2
by Fittkau (1973) this would be the equivalent of 80-120 kg/km 2 of nutrient rich
faecal material entering the system daily. Such quantities are apparently ade-
quate to increase phytoplankton (Zaret et al. 1981) and fish production (Fittkau
1973). Another aspect of their ecological importance is that as a top predator,
there are reports of increases in piranha (Serrasalmus spp.) and capivara popula-
tions in areas whcrc thc cairnans have been seriously rcduced (Plotkin et al., in
press; Medern 1981).
The rernaining genus of caiman includes the dwarf and srnooth-fronted cairnans
(Paleosuchus palpebrosus and P. trigonatus) (Photo 12) which are the smallest of

Table JO Predators 01 halchlin12 ('({{II[({II <ro"o"il//,\,

Rcptilia :\ I'CS

Eunecles muritllls An/ca (ocui

Cuimun crocodi/l/,\ 1"hiru mvcleria
MelanoslIchus /liga ,V/I'Cleria albus
Mammalia Anhinga al/hingu
Racoons (ProcYOI/)
Foxes (Ceri!ocvolI!
Jaguars (Fc/is)

References: Staton & DiXO!1 197'i

402

I'holo 12 The dwarf caimun I'all'uslichus is still common in the small forest streams of Am3zonia.
(Photo: William Magnus,on)

the caimans (maxImum length P. trigonatus 2.26 m; P. palpebrosus 1.55 m

(Medern 1981). These caimans are typical of forest streams and small rivers with
fast currents and are usually solitary (Medern 1960, 1967, 1971, 1981; Campbell
1973; Oixon & Soini 1977; Oonoso Barros 1966; Magnusson 1980; Godshalk
1982). The two species may be sympatric through much of thcir range although
where this occurs, one species is usually predominant (Goldshalk 1982; Medern
1967, 1971, 1981) and hybridization may occur both in the wild and in captivity
(Medern 1981). These species may be found together with Caiman; however, they
are gene rally much less abundant and tend to be in more protected sites such as
under fallen trees or in srnall bushes. P. trigonatus does not bask in open sunlight
whereas P. pa/pehrosus cornrnonly does, possibly indicating that the former
species is more adapted to the deeper forest streams where the thicker canopy
does not allow penctratlon of thc sun's rays (Medern 1971). Natural predators of
Paleosuchus are the anaconda (Eunectes) , jaguars and smaller spotted cats, and
other caimans. particularly Caiman which may overlap in niche distribution. This
species does not have d comrnercially valuable skin, however their meat is
commonly eaten in Peru (Medern 1981) and is traded illegally trom the upper
Brazilian Amazon tu both Peru and Colombia (Best, unpublished).
They feed on a tyrically wide variety of iterns such as snails , crabs, fish, frogs
and srnall marnmais (Table 11). Gastroliths are common in both species (Van-
zolini & Gornes 19 7 9; Medern 1967. 1981). The reproductive biology of Pa-
leosuchus is pOOfl)' k nown. ami its egg was first described by Oonoso-Barras
 403

Tahle 11 Prey species of the smooth-fronted and dwarf caimans, PaleoslIchus palpehrosus and P.
trigonatus.

Inseto Amphibia
Dytiscidae ()sleocephalus taurinus
Orthoptera l.cplOdactylus sp.
Colcoptera Pisces
Diptera Serrasalmus sp.
Arachnida Cichlidae
Spiders Engraulidac
Crustacca Hoplias malaharicus
Crab ('1 Trichodactvlu.1 peruviunus) Aves
Shrimp Ikron (Ardeidac)
Mollusca C\'(/nocorax violaceus
Pomacea sp. Mammalia
Reptilia Myoproela sp.
Caiman crocodilus Rodent
Paleosuchus sp. Marsupial
Snakc

Refercnces: Medern 1981: Vanzolini & Gomcs 197,).

(1966). Since that time a number of nests have been described and a hybridization
in captivity has been reported. Paleosuchus nests are usually found within 3-5 m
of the sm all streams where these species occur (Medern 1981; Dixon & Soini
1977). The nest is composed of earth, leaf-litter, twigs and other debris and may
attain a size of 120-146 in diameter and 45-49 cm in height. The temperature
inside the nest may be some 4° C above that of the environment (Medern 1971,
1981; Dixon & Soini 1977). Clutch sizes are between 10-13 eggs in P. trigonatus
and 13-18 eggs in P. palpebrosus (Medern 1971, 1972, 1981; Dixon & Soini 1977).
The incubation period was about 126 days for the P. trigonatus X P. palpebrosus
hybrid in captivity (Medern 1981). There is some doubt as to whether the female
guards the nest as do other cairnans. Newly hatched Paleosuchus do not go
directly to the water, but spend at least two days hidden near the nest while a
mucous layer over their skin drics up (Medern 1972). Very young Paleosuchus
have an ochre-coloured head and are often called jacare de cara encarnada or red-
faced caimans by the locals because of this. Recent hatchings are difficult to find
near other size-classes of Paleosuchus or Caiman and may seek shallow, srnall
creeks or pools to avoid inter- or intraspecific predation (Magnusson 1980;
Carnpbell 1973).

Summary

The biology and ecology of the aquatic mammals: the Amazonian manatee
404

(Trichechus inunguis) , giant otter (Pteronura brasiliensis) , river otter (Lutra

enudris) , capybara (Hydrochoerus hydrochaeris) , freshwater dolphins (Inia
geoffrensis and Sotaliafiliviatilis), as weil as the aquatic reptiles: turtles (Podoc-
nemis, Peltocephalus, Chelus, Phrynops, Rhinoclemmys, Kinosternon, Plate-
mys), snakes (Helicops, Hydrodynastes, Hydrops, Micrurus and Eunectes), liz-
ards (Neustricurus, Crocodilurus and Dracaena), and caimans (Melanosuchus,
Caiman and Paleosuchus) is reviewed with regard to their ecological inter-
relationships within the Amazonian aquatic ecosystem and their past and present
status in relation to human exploitation.

Acknowledgements

I thank Professor H. Sioli and Dr. W.E. Kerr for the stimulus they have given me
and many other researchers to study in the Amazon. I also thank Dr. R.
Shrimpton, Vcra M. F. da Silva for reading the manuscript and especially George
Nakamura who meticulously corrected and typed the manuscript. Dr. W.E.
Magnusson provided some of the critical literature on reptiles. The Aquatic
Mammal Division of INPA is supported by thc National Research Council of
Brasil (CNPq). with additional funding from ELETROBRAS/CELPA/ELE-
TRONORTE, the World Wildlife Fund (Int/US), the International Union for the
Conservation of Nature and National Resources (IUCN), the Secretaria Especial
do Meio Ambiente (SEMA), the Ford Foundation (Brasil), IBM (Brasil),
Governo do Estado do Amazonas, the Vancouver Public Aquarium and The
Cousteau Societv.

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