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3.10 Carnivores

3.10.1 Pteronura brasiliensis (giant otter)
3.10.2 Felis pardalis (Ocelot)
3.10.3 Panthera onca (Jaguar)

Latin American wildlife numbers 66 carnivore species (excluding seals), including 14 canids, three bears, 16 Procyonidae, 22 mustelids, and 11 felids (285). Carnivores have traditionally been culled (justifiably or not), hunted for sport, or commercially for their skins, or, to a lesser extent for food, e.g. Nasua and Potos.

The main fur species are the foxes (Dusicyon culpaeus, D. griseus, D. gymnocercus, D. thous) in southern South America (91, 362, 451, 489, Mones, pers. com.; Rottmann, pers. com.), the aquatic mustelids (Lutra spp. and Pteronura brasiliensis) and the felids Felis pardalis. Panthera onca, and, to a lesser extent, F. wiedii and F. concolor in the tropics and F. geoffroyi and F. colocolo in the southern part of the continent (44, 105, 244, 380, 421, 451, 545).

The IUCN Mammal Red Data Book (576) lists Ursus arctos nelsoni of northern Mexico as an endangered species. Canis lupus, Chrysocyon brachyurus, Speothos venaticus, Tremarctos ornatus, Lutra felina, L. platensis (which is identical to L. l. longicaudis, according to Zyll de Jong, 637), Pteronura brasiliensis. Felis pardalis, F. tigrina, F. wiedii and Panthera onca are all listed as vulnerable. Felis jacobita is listed as rare whereas Lutra provocax and Felis yagouaroundi are of indeterminate status, and Atelocynus microtis is listed as insufficiently known. There are serious problems of conservation and management for a number of the more valuable fur species and this is particularly true of Pteronura brasiliensis, Felis pardalis and Panthera onca.

3.10.1 Pteronura brasiliensis (giant otter)

Vernacular names: Ariraí (Argentina, Bolivia, Paraguay), ariranha (Brazil), lobo corbata (Uruguay), lobo de río (Peru, Bolivia), lobo de río grande (Argentina, Paraguay), perro de agua (Colombia, Venezuela), water dog (Guyana), watradagoe (Suriname).

Geographical variation and distribution: Pteronura brasiliensis is strictly South American: the typical subspecies P. b. brasiliensis has its distribution in the Orinoco, Amazon and Guianas river systems, whereas the southern P. brasiliensis paranensis ranges from southern Brazil, Paraguay and Uruguay to the northeastern tip of Argentina (91). Local extinctions have reduced its distribution, which is now discontinuous (122, 245, 366, 398, 576).

Elevational range: Along tropical rivers from sea level (168, 245, 398).

Size and weight: This is the world's largest mustelid. It exhibits sexual dimorphism by size: the total length of adult males ranges from 150 to 180 cm (and reportedly up to 220 cm), a third of which is the tail, and it weighs 26-35 kg. Adult females run from 150 to 170 cm and 22 to 26 kg (113, 168, 398).

Habitat: P. brasiliensis is an amphibious species occupying a great variety of bodies of water such as slow-flowing rivers and creeks, lagoons, swampy areas, swamp forest and flooded forest. In Suriname, and possibly in other areas with similar seasonality, fish availability seems to be decisive in habitat selection. Giant otters concentrate in creeks in the dry season and disperse into flooded forests in the rainy season. They prefer shallow, blackwater creeks and avoid brackish waters and mountain streams. One requirement is a dry, tree-shaded spot along the shore on which to build their dens and campsites (78, 168, 245, 328).

Abundance: Originally, this was apparently a common species in its preferred habitat. The maximum known population density is 1.2 animals per km in the Suriname creek according to Duplaix (168). Laider (328) reports one otter per 6.5 km in Demerara, Guyana. The species is very rare today throughout its range, however, except for certain protected areas (576).

Behaviour: This diurnal animal lives in stable family groups, usually made up of a mated pair and their cubs: subadults may remain in the parental group up to the age of two. While group size may range from 2 to 20, 3 to 8 is more common. The diurnal and gregarious habits of the giant otter, its complex vocal communications, its territorial calls and its trusting nature in the wild make it conspicuously easy to detect. Its gait is clumsy on land but it moves with extreme ease in the water (78, 168, 328, 398, 495). Duplaix (168) describes some of the fascinating aspects of her research on the sociobiology of the species.

Feeding habits: Giant otter are fish-eaters, eating mainly medium-sized (15-30 cm) bottom fish in brackish waters, and sometimes rounding out the diet with crabs. They customarily fish in groups, which apparently increases efficiency, but each otter eats its own catch (78, 168, 328, 398). Hoplias malabaricus made up half the catch in Suriname (168). They sometimes fish in deep water in mid-river but shore fishing is more efficient (328). Each individual consumes an estimated 2.8-4.0 kg of fish every day (168).

Reproduction: It is not known at what age giant otters reach sexual maturity but probably after the age of three, as Duplaix (168) reports that subadults disperse at the age of two to three years and then form a pair bond. They apparently give birth once a year during the low-water period (August-October in Suriname) in their dry-season home areas (168). The gestation period is 65-70 days. While litter size varies from one to five, two to three is more common (29, 168, 495, 583). The neonates are quite undeveloped, weighing some 200 g, and open their eyes only 30 days after birth. At the age of three months the cubs begin to eat fish (29, 168).

Hunting: Little documentation of hunting methods is given in the bibliography. Shooting giant otters in the water appears to be the usual (but wasteful) method, various authors (122, 328, 398) reporting that a good many animals shot in this way simply sink and are not retrieved. Massoia (366) mentions the use of snares. Hunters in the llanos region locate inhabited dens during the dry season and the otters are clubbed to death as they emerge. The hunters thus obtain a pelt free of bullet-holes (Ojasti, pers. obs.). The natives in Guyana club sleeping otters to death at night in their campsites (327).

Products: The giant otter pelt has very short but dense smooth, bright, chocolate-coloured fur and is highly prized on world fur markets. Referring to this pelt, Dourojeanni (163) cites a European import price of US$90 in 1970: only jaguar skins were worth more. The otter is skinned by a ventral cut from the mouth to the tip of the tail, and the open skin is stretched out to dry, held down by wooden pegs or nails.

P. brasiliensis were already moderately hunted in the last century in Brazil to make purportedly waterproof coats (89). More recent statistics from 1959-1969 for the Brazilian Amazon (105, 545) report annual export figures of 1 000-3 000 pelts, gradually declining to a few hundred and then to only 12 pelts in 1971. No statistics are available from other countries of the area, but this species has probably been commercially hunted very intensively throughout its entire range.

Management: Intense commercial hunting in recent decades, spurred by the high unit price for giant otters, has drastically reduced P. brasiliensis populations. The IUCN considers the species to be vulnerable (576). It is the most endangered aquatic mustelid in the world (328), listed on Appendix I of CITES and explicitly protected by law at least in Argentina, Brazil, Colombia, Guyana, Peru, Suriname and Venezuela. The following traits make it particularly vulnerable to hunting: 1) size, diurnal habit, gregarious behaviour and lack of suspicion, 2) fragile and complex social organization, 3) late sexual maturity, and 4) habitat (river networks are the main channels of communication and travel in forest regions).

Restrictions implemented by CITES member countries since 1973 on the international skin trade are thought to have substantially reduced commercial hunting of giant otters, and they may be slowly recovering in some areas. Extensive deforestation, sedimentation, hydraulic changes in rivers and pollution may constitute new threats for the species in some regions (168). As a top carnivore, P. brasiliensis would obviously be highly susceptible to chemical pollution (168).

Clearly, the top management priority in the immediate future must be strict enforcement of the protective legislation already on the books to ensure the recovery of wild populations (380). Captive breeding, suggested by a number of outstanding authors (122, 432, 576, 582), presents serious limitations as an effective conservation tool in that: 1) it requires the initial capture of wild populations, 2) social intolerance in captivity often means mortality from fighting, 3) successful breeding in some zoos (Caracas, Paramaribo, Sao Paulo) is offset by the fact that most captive adults have killed their cubs, 4) large-scale captive breeding is extremely expensive because of the animal's size, diet and the type of installations required (captive giant otters are highly dangerous to humans) (29, 138, 168, 582).

3.10.2 Felis pardalis (Ocelot)

Vernacular names: Cunaguaro (Venezuela), gato onza (Argentina), gato tigre (Panama), heitigrikati (Suriname), jaguatirica (Brazil), manigordo (Costa Rica, Panama, Venezuela), maracajá (Brazil), ocelot (Belize, Guyana, Suriname), ocelote (Argentina, Colombia, Costa Rica, Paraguay, Peru, Mexico, etc.), tiger cat (Belize), tigrecillo (Bolivia), tigrillo (Colombia, Ecuador, Guatemala, Peru).

Geographical variation and distribution: From Arizona and Texas in North America (excepting the Mexican altiplano) down through Central and South America to northern Peru on the Pacific slope and Tucumán, Argentina on the southeastern side (323, 380). Five subspecies are recognized in Central America (253) and 5 more in South America (91).

Elevational range: From sea level, concentrated in the tropical belt up to roughly 1 000 m, but occasionally reported at higher altitudes (66, 245, 382, 404).

Size and weight: Total adult length varies from 95 to 135 cm, a third of which is the tail. An ocelot stands some 45 cm high at the shoulder and weighs from 7-14 kg, depending on the subspecies and sex: the males are usually rather more corpulent than the females (294, 336, 348, 353, 382).

Habitat: F. pardalis occupies a great variety of habitats, prefers wooded areas ranging from mangrove swamps and tropical moist forest to dry forests, scrub forests and stubble, but may also be found on savannahs, beaches and other open areas (17, 66, 134, 141, 181, 336, 348, 382). In the llanos its preferred habitat is the gallery forest but it may roam widely into the savannah at night (353, 406).

Abundance: As a top carnivore the ocelot is relatively scarce. The population density in Venezuela in a protected area of gallery forest, low forest and savannah is estimated at 0.25/km2 (174) and 0.40 (353). Smith (544) estimates one ocelot per 10 km2 in Amazonia, and Emmons (181) reports 0.8/km2 in the Peruvian Amazon.

Behaviour: The ocelot is sedentary, solitary and secretive. Adults occupy a permanent home range of 76-930 ha (averaging 346 ha) according to Ludlow's summary (353). Males have an exclusive and large home range overlapping that of the females but animals sharing the same range consort only to mate although some authors do postulate that ocelots live in pairs. Ocelots are much more active at night: diurnal activity is variable and depends on the habits of its main prey - like most felids, ocelots stalk their prey. They are good climbers and often rest in the branches of trees but most of their activities take place on the ground (17, 181, 182, 353, 382. 406, 572).

Feeding habits: The ocelot is a carnivore par excellence, feeding primarily on mammals. Its habit of using permanent latrines facilitates studies of ocelot diets. The most frequent quarry in one savannah forest mosaic in the Venezuelan llanos consisted of small rodents (Zygodontomys, Sigmomys, Holochilus), crabs (Dilocarcinus dentatus), iguanas and rabbits (353). In forest areas, rodents (mainly Oryuzomys) and small marsupials (Marmosa), the Cayenne spiny rat (Proechimys), agoutis (Dasyprocta) and other medium-sized mammals (Didelphis, Sylvilagus, Agouti, Dasypus, Nasua) make up the basic diet, supplemented by birds, particularly tinamous, reptiles, fish, crabs and the odd insect (57, 181, 182, 294, 336, 406, 572). The frequency of the various quarry in the ocelot's diet parallels prey abundance in the habitat, Emmons (181) reports, as this felid is apparently fairly unselective. The same author estimates the daily consumption at 558-837 g. This cat may occasionally attack poultry and the young of domestic animals.

Reproduction: Sexual maturity probably occurs between the ages of two and three with the animals mating once a year (or at most, according to Eaton (170) every nine months), usually between September and April, according to the available data (170, 182, 336, 353, 382). The male is believed to mate with females (usually three in number) whose home range overlaps his own (181, 353). The gestation period of about 80 days produces a litter of one to three cubs, averaging 1.3 (170). The cubs require maternal care for many months and may remain in the mother's territory as long as two years before moving on (170, 535).

Hunting: Professional hunters work mainly with traps made from tree trunks or snares baited with monkeys or wild birds. They also hunt with dogs that tree the ocelot, whereupon the animal can easily be dispatched with a shot. Or they may hunt at night with lanterns (66, 258, 336, 544). Hunting expeditions may involve journeys of hundreds of kilometres and last for several months. Many kills, however, are the product of a casual encounter between an ocelot and a campesino with a gun.

Products: The fur of the ocelot is plush, bright and very strong, patterned with spots and patches on an ochre-yellow to orange-yellow background and highly prized in the industrialized countries, mainly for ladies' coats, which has spurred heavy commercial hunting for export. The pelt is prepared by a ventral cut from the mouth to the tip of the tail, and then to the tips of the paws. The skin is dried in the shade, with or without salt, stretched and held with pegs or nails, and sprayed with insecticide to prevent pest attacks. The meat is sometimes eaten and the easily tamed cubs are in great demand as pets (US prices can be as high as US$800 according to the IUCN figures of 1982) (66, 170, 323, 336,421, 544).

Ocelot skins were the top seller among neotropical mammal skins (Table 16). Although the available statistics (44, 97, 105, 245, 334, 380, 421, 479, 544) are fragmentary and expressed in different units (numbers, weight or value of skins), and sometimes more than one species is included under a single heading, it is easy to hazard a guess that during the apogee of commercial felid hunting in the 1960s, some 200 000 ocelot skins were exported legally every year. During that same period the US imported roughly 130 000 skins a year, with appreciable quantities also being imported by the Federal Republic of Germany, England and other European countries. Legal restrictions, and possibly the decline of fiercely exploited ocelot populations, drove the trade down in the 1970s to an annual 10 000-40 000 skins, and the main trade axes shifted from Brazil/US to Paraguay/Federal Republic of Germany (26, 576).

Commercial hunting of spotted cats offered employment to thousands of "gateiros" or cat hunters, occasional income for a great many campesinos, raw material for the fur industry and a source of hard currency. This species is therefore a potential candidate for rational utilization of wildlife resources.

Management: Commercial ocelot hunting may well have far exceeded recommended sustainable harvest levels for populations in areas accessible to hunters, thus reducing F. pardalis distribution, abundance and productivity. Additionally, though ocelots do tolerate partially modified habitats, they may be seriously affected by extensive deforestation (245, 323, 353, 404, 421, 544). The IUCN considers ocelots to be vulnerable (576). CITES lists them in Appendix 2 (the subspecies F. pardalis mearnsi of Central America and F.p. mitis of southeastern Brazil, Paraguay and northern Argentina are listed in Appendix 1), and ocelot hunting and trade is prohibited in all countries of Latin America.

Calling a halt to indiscriminate ocelot hunting, already partially successful, is the key to managing this valuable species and should allow ocelot populations to gradually recover. Future management should be based on reliable population and productivity data for different habitats as a basis for setting sustained harvest criteria. The harvest could never be very high, however, given the ocelot's late sexual maturity and low reproductive capacity. As soon as populations had expanded sufficiently, administrative mechanisms would need to be designed to allow producer countries to harvest on a sustained base. This should be done experimentally, with built-in guarantees for maximum economic benefits to rural people and systematic monitoring to ensure a sound basis for the final legislation. It would also be important to consolidate the coordination of international trade in accordance with CITES guidelines.

Captive breeding: Ocelots adapt well to captivity and are found in a number of zoos. The record longevity in captivity is 18 years (138, 170, 310). Coimbra Filjo (122) and Koford (323) suggest captive breeding as a strategy. Some zoos have successfully bred ocelots, even achieving two litters in one year with high survival provided the neonates are separated immediately after birth, but the supply of animals born in captivity is generally insufficient (138), 170) and wild inputs have to be brought in to maintain exhibitions.

3.10.3 Panthera onca (Jaguar)

Vernacular names: Jaguar, yaguar (widely used), onca pintada (Brazil), otorongo (Peru), pakira tigri (Suriname), tigre (widely used), yaguareté (Argentina).

Geographical variation and distribution: Once ranging from Arizona and Texas in the United States to central Argentina in the eastern Andes, the jaguar is now extinct in the US, El Salvador, Uruguay, the Argentinian Pampa and many other populated parts of Latin America. Eight subspecies are recognized. P. onca is also found in the literature as Felis or Leo onca (91, 195, 253, 478).

Elevational range: Jaguars are usually found in the tropics up to 1 000 m, though occasionally as high as 3 000 m (17, 195, 245, 348, 406, 576).

Size and weight: P. onca is the largest American felid: size depends on sexual dimorphism and geographic variation. The total length of adult males in Mexico varies from 160-225 cm and the weight from 64-114 kg, and for females 140-185 cm and 45-82 kg (336). The reported figures for P. onca palustris in southern Brazil and adjoining areas are 195-240 cm and 80-120 kg for males and 185-200 cm and 60-90 kg for females. The top weight is 130 kg on an empty stomach and 150 kg on a full stomach. The tail is 50-70 cm long and the shoulder height 60-100 cm (14, 15, 195, 248, 348, 382, 406).

Habitat: P. onca prefers the tangled moist forest along rivers, swamp and mangrove forest but is also found in deciduous forest, semi-arid and mountainous zones. Jaguars avoid open savannah and tend to shun heavily modified habitats. Prey abundance is clearly a decisive factor in habitat quality (14, 17, 195, 336, 382, 405, 487, 521).

Abundance: Jaguar abundance has been estimated by reading jaguar tracks, by radio-tracking and by the reported kill ratio within a given area over a short period of time. The maximum densities recorded in optimum habitat were roughly 1 jaguar per 13 km2 in a reserve in Belize (487) and 1 per 25 km2 in Pantanal de Mato Grosso, Brazil (15, 521). Eisemberg et. al. (174) estimate 1 per 50 km2 and Crespo (141) 1 per 55 km2 in the national parks of Guatopo in Venezuela and Iguazú in Argentina, respectively. Smith (544) gives an average of 100 km2 per individual in the Brazilian Amazon. Recent estimates (301) extrapolated from average densities over large areas give a total figure of 150-200 jaguars for Costa Rica, 4-5 000 for Venezuela and 1 800 to 3 500 for the Pantanal de Mato Grosso (301).

Behaviour: The jaguar is primarily a nocturnal and crepuscular, solitary, and sedentary animal. The females occupy partially overlapping home ranges, which they share with their subadult cubs, of 25-38 km2 according to the available data (487, 521). Male home ranges are far larger and may overlap those of two or more females, but not the territory of other adult males. They mark their territories by loud roaring, by urine, and by scratch marks on trunks. With the exception of females in oestrus, or with cubs, they seem to lead a solitary existence. They may travel kilometres every night or they may remain for two or three days near a recent kill. During the hot hours of the day jaguars like to rest in the thick vegetation or occasionally along branches up a tree. They frequent swampy areas and beaches and swim often and easily (14, 17, 181, 195, 248, 336, 406, 521).

Feeding habits: Jaguars eat a great variety of vertebrates: rodents, rabbits, armadillos, sloths, ant-bears, primates, peccaries, capybaras, deer, large birds, iguanas, snakes, caimans, water turtles and tortoises and their eggs, and fish too. Emmons (181) estimates the average weight of their Peruvian Amazon prey at 7.2 kg and their daily consumption of meat at 1.4 kg (for animals with an average weight of 34 kg). In inhabited areas they may prey upon domestic animals - pigs, dogs, horses and even full-grown cattle, and can occasion quite considerable losses. It is believed that only a few specimens are in the habit of eating livestock whereas most feed in the wild (14, 17, 195, 294, 336, 348, 379, 406, 487). Jaguars usually ambush their quarry at short distances, killing the larger ones by leaping on them and inflicting a mortal neck bite. The prey is then dragged off to a tree-shaded place (14, 521, 524).

Reproduction: Jaguars reach sexual maturity in three years and are polyoestrous with a 6-17 day period of heat (138, 529). They apparently mate once a year at no particular season, at least in tropical areas (17, 348, 382, 406, 529) giving birth to a litter of one to four cubs (an average 1.8 cubs in captivity). The usual figure is two cubs with a birthweight of 0.8-0.9 kg. They are cared for by the female for several months and remain in the mother's territory up to the age of two (267, 336, 382, 421, 529). The record longevity, in captivity, was 23 years (138).

Hunting: Hunting with dogs is the method of choice: the dogs find and give chase to the jaguar, which sooner or later will opt to climb a tree or turn at bay on the ground, whereupon the hunters come up and shoot it (in former times jaguars were speared). Hunters often hide at night near a tree baited with a half-eaten carcass or other dead or live bait, lure the jaguar by imitating its roar, or hunt at night from canoes. Some professional hunters also trap jaguars, but this is fairly ineffective compared to ocelot trapping. Stock-owners sometimes combat stock-eating jaguars by putting out poison bait. Aside from intentional hunting, many jaguars are killed as the result of casual encounters. Jaguar is also a prized target of certain sport hunters. In livestock regions jaguars may be culled to protect the herds (14, 17, 66, 94, 336, 379, 382, 406, 544).

Products: The jaguar's ample and colourful pelt has been a spur to commercial hunters since the early 1800s (195, 234, 451), earning the top unit value for neotropical furs, amounting to some 100-300 US dollars for the hunter (232, 544), and ranking in third place for export value (Table 16).

The history of jaguar exploitation parallels the ocelot's, with a sharp upward swing in the 1960s. In the late 1960s between 7 758 and 13 516 skins were exported (421). The annual kill in Brazil topped 15 000 (544). Then protectionist laws were passed and jaguar hunting and the skin trade were curtailed.

Management: The triple pressures of commercial hunting, sport hunting and (until recently) culling, combined with growing encroachment on its habitat, has drastically reduced P. onca populations. Today the jaguar is considered vulnerable (576), listed on CITES Appendix 1, and hunting is totally prohibited in all countries of the area excepting Mexico and Belize, which have established a limited quota of licences for sport hunting, and Costa Rica and Peru, where hunting may be authorized for individual specimens preying on livestock (301). Species abundance figures for some areas are approximate, and despite rather intensive poaching, some populations may be on the road to recovery, whereas others are endangered or have become extinct.

A recent symposium (301) raised the option of opening limited, very expensive hunting licence quotas, on the Mexican pattern, to generate funds for livestock research and conservation, as opposed to tolerating poaching. However, outstanding experts (139, 380) feel it would be counter-productive to reopen hunting, as the jaguar situation is extremely fragile. They propose focusing research efforts on the current status and productivity of jaguar populations in order to provide guidelines for conservation and future management.

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