The idea of writing a guide to Central American carnivores came to us after spending many years working as researchers and conservationists in the Neotropics. Together we have spent over 13 years involved in several aspects of tropical biology, research, education, and conservation, mostly in Costa Rica and Nicaragua. In recent years, we have come to realize that the status of many species of tropical carnivores is critical. Some of the largest and most recognizable species, like the jaguar and the puma, have suffered tremendous losses to their habitats. Some of these losses have become the main reason why these species are doomed to disappear entirely from many parts of their ranges. While conservation efforts in Central America are progressing steadily throughout the region, for some of these species, the progress simply is not happening fast enough.
In this book we make some sobering observations and claims as to the future of several species of carnivores in the region. These observations are based on the present knowledge of the biological needs of the species and the immediate prospects for habitat conservation and restoration. While the situation is not terribly urgent for many species of Neotropical carnivores, for several of them the next 10 or 15 years will decide their future presence and sustainability or their disappearance from the region.
Regional efforts, such as the Central American Biological Corridor, or Paseo Pantera, are initiatives in the right direction, although the difficulties in establishing multipurpose biological corridors through an assortment of countries, economies, and cultures are numerous and complex. Conservationists tend to become involved in politics, sociology, and economics, often to the point of losing sight of what is actually best for the species and habitats they are trying to preserve. If we really want to make progress in the conservation of carnivores and their habitats, biological information about the species should be made widely available to all segments of society, not just to biologists. Politicians, economists, sociologists, and conservationists need to speak the same language. Conservation in the American tropics involves much more than choosing where parks, preserves, and biological corridors should be established. It also requires striking a balance among the specific needs of a dozen countries (some outside Central America, but with strong influences in the region) and taking into account their economic and social pressures, as well as incorporating the needs of future generations in the decision-making process.
We hope that the information in this guide, and in similar guides to come, helps bring relevant biological information to the discussion and negotiation table. Rarely is the biologist the one who decides where a conservation area will be located, nor is it the biologist who writes the laws or attempts to enforce them. This guide, then, is written with the nonbiologist in mind. Most technical terms are either explained in the text when first used or included in the glossary.
How to Use This Book
This guide is divided into three parts. The introductory chapter deals with general characteristics of Central American carnivores, including their evolution, taxonomy and classification, distribution, habits, and other topics, such as vegetation types and tracking. Following the introduction, each family of Central American carnivore is presented, beginning with an introduction for each family and followed by a description of each species. We include with each description a map of species distribution where the reader can find the areas most likely to have representative populations of the species. These distribution maps differ from those commonly found in most field guides in that, in addition to showing the historic range for the species, we include the major patches of suitable habitats where the species actually can be found at present. In general terms, these patches correspond to protected areas, both private and government-supported (for example, national parks), or unprotected areas with various types of natural vegetation cover. We make no attempt to qualify these protected areas as suitable or unsuitable for specific species; as we will show in the species accounts, there is much to be learned about the habitat requirements, territories, and home ranges for most species, and this kind of information--which only long-term research can provide--is essential for managing these areas and maintaining viable populations of each species. At the end of each species description there is a conservation status summary of the species intended for quick reference. Finally, a chapter on carnivore conservation explores some of the major carnivore conservation topics and presents other pressing issues relating to the future of carnivores in Central America.
We have strived to include the latest research findings and taxonomic changes in this guide. However, carnivore taxonomy and systematics, as well as ecological research, is a very active and continuously changing field, and not all of the players are in agreement with the findings of their colleagues. We have included in the species accounts only that information which has been published in peer-reviewed scientific journals, books from recognized and reviewed publishing houses, and reports and publications from agencies involved in carnivore conservation, such as the International Union for the Conservation of Nature (IUCN), the World Wildlife Fund (WWF), and others. Our firsthand accounts of encounters with individuals of most species are intended to give insights into the lives and habits of these creatures, the places where they live, and some of the risks to which they are exposed in Central America. If these descriptions succeed in raising readers' interest in knowing a little more about these species, our goal will have been fulfilled.
It was about 11 p.m. on a moonless night in Guanacaste, northern Costa Rica. In the enveloping darkness, Manuel and Javier, caretakers of the Maritza Biological Field Station in Guanacaste National Park, returned on horseback from a trip to the nearest town some io miles away. Loaded with supplies and tired of the long ride, they allowed their horses to pick their footing on the narrow trail that led to the station through the thick forests that blanketed the sides of the Cacao and Orosí volcanoes. Their flashlights were tucked away in their saddle bags. They trusted their mounts to follow the familiar path. A light rain filtered through the canopy and drenched the exposed soil, creating a sticky mud that made sucking noises at every step of the horses. There was near total darkness. Horses are known to have excellent sense of smell, and both horses kept their noses down, perhaps following their own scent which was left there on their previous passing.
Suddenly, Javier's horse, who was leading, stopped and snorted nervously. His ears pricked forward toward the blackness ahead. Manuel, some 50 yards behind, spurred his own horse to keep it going, finding it difficult to make it obey his orders. When it reached Javier's, both horses refused to continue, snorting and trying to turn back on the trail.
"I bet there's a snake on the trail," Javier said, reaching back to his saddle bag for the flashlight. Manuel followed suit. They both turned on their lights and the bright beams probed the darkness, illuminating the trail in front of them. They could see nothing. The horses still refused to go on, requiring the riders to keep tight reins lest they lose control of their mounts. A heavy feeling of foreboding began to fill the men's hearts. With their lights they scanned the trail and the forest directly ahead of them, searching for the cause of the horses' near-panicky behavior.
"Oh my God!" Javier whispered between clenched teeth. Manuel's light beam joined Javier's. Reflecting some 50 yards ahead on the trail were the two incandescent coals of the eyes of an enormous jaguar, framed by a round face about the size of a soccer ball. It was lying in the middle of the trail, motionless, attention fully concentrated on the intruders to its territory. Time seemed to stop. For several interminable minutes the jaguar and the men awaited one or the other's next move...
Jaguars are not commonly seen in the wild. Recently, in Costa Rica's Santa Rosa National Park, there have been several daytime sightings of jaguars, including a female with a cub that boldly strolled into the park's camping area in the middle of the day, creating a general panic from the campers that resembled a five-alarm fire drill at an elementary school. Santa Rosa's jaguars, having enjoyed almost 20 years of continued protection, have lost some of their long-time acquired fear of people and have been sighted along beaches and rivers where they feed on turtles and other small animals. In the forests that cover the volcanoes, jaguars are more secretive and nocturnal, although recently there had been several recorded attacks on horses that were kept at the biological station.
"What do we do now?" Javier asked, his voice quiet but full of consternation. The men's spirits grew cold from dread and the chill of the falling rain. As long as the jaguar stayed on the trail it would be impossible to go through, and the horses were not about to move an inch forward. As if reading their thoughts, the jaguar stood up and, with a fluid and soundless move, disappeared into the thick vegetation by the side of the trail. There was neither the sound of branches breaking nor of leaves being crushed. As far as they knew, the jaguar could be crouching by the trail, waiting for either of them to approach. If this was the same jaguar that was killing the station's horses, it might sense a familiar smell that it associated with food.
The men turned off their flashlights. The surrounding darkness was made even darker by their blindness following the intense light. They could hear the sounds of their own hearts pumping wildly in their chests. Could the jaguar hear them, too?
The horses began to walk cautiously, their noses close to the ground. The men turned their lights back on and scanned the trail. At the place where the jaguar had entered the forest, there were tracks larger than the closed fist of a man reflected in the lights. They could be seen along the trail, a witness to the jaguar's fondness for using the open paths made by human and beast. The horses picked up the pace without any prodding from the riders. Of the jaguar, only the indelible image remained--an image that for weeks and months to come fueled stories of the men's close encounter with the most formidable predator that roams the forests of Central America. --C. De la R.
Taxonomy and Relatives
One of the first written references to the American jaguar comes from Amerigo Vespucci in 1500, who mentions "panthers" as one of the animals present in Venezuela (Hoogesteijn and Mondolfi, 1992). The name jaguar comes from the Tupiguaraní language (a widespread language in the Amazon and other parts of South America), which translated means "wild beast that dominates its prey in one jump." Yaguareté, another common indigenous name still in use by the Guaraní Indians of Paraguay, means "body like a dog." The species was formally described by Linnaeus in 1758, who used the common name "onza" as part of the scientific name (Panthera onca). Its closest relatives in Central America are the puma (Puma concolor), the ocelot (Leopardus pardalis), the margay (Leopardus wiedii), the tiger cat (Leopardus tigrinus), and the jaguarundi (Herpailurus yaguarondi). Its closest taxonomic relatives are the African lion (Panthera leo) and the Indian tiger (Panthera tigris). The jaguar is the third largest cat species in the world. To date, up to eight subspecies have been recognized. Recent taxonomic revisions have split the genus Felis into four genera, Panthera, Leopardus, Felis, and Herpailurus.
Jaguar, tigre, tigre real, yaguar (Spanish); zac-bolay (Mayan); onça negra, yaguara pichuna (Brazil); yagua-hu, tigre negro (black jaguar), American tiger; onza (Brazil, Venezuela); yaguareté (Argentina, Paraguay).
There is an interesting theory that explains the massive size of the jaguar. Compared to tigers and lions, the jaguar is very similarly equipped in terms of weapons (teeth and claws) as well as raw power (strength). In continents such as Africa where there are many herding species of large prey such as antelopes, zebras, and buffaloes (common prey of lions), it is easy to explain the need to develop massive musculature and formidable weapons for overpowering them. In Central and South America, however, there are few large animals that might require such strength (tapirs and, perhaps, deer). Most of the present-day natural prey of the jaguar consists of much smaller animals such as peccaries, brocket deer, capybaras (in South America only), armadillos, agoutis, and other even smaller game, few of which travel in herds. According to the theory, when jaguars, pumas, and their extinct relative the saber-toothed cat arrived from North America during the glaciation periods, they encountered a fauna quite different from the one we have today. Gomphotheres, giant sloths, giant armadillos, mastodons, and even a species of rhinoceros populated the forests and savannas of Central America. These animals became extinct relatively recently (according to one theory, in part due to overhunting by early humans).
It is in recent times that other large prey, suitable for the jaguar's strong weaponry, arrived in the region: horses and cattle introduced by the Spaniards during the conquest of the Americas. Interestingly, jaguars are unique among the big cats in that they are the only ones that regularly kill their prey by piercing their skulls, while other species tend to prefer suffocation or strangulation methods.
Jaguars resemble the African leopards in build and coloration, although they are more robust, the head and front paws being larger and the tail shorter. In relation to its body size, it is the strongest of all cats and is able to overpower and move prey heavier and larger than itself. We observed the remains of an adult horse in northern Costa Rica that was killed by a jaguar at night in an open pasture and dragged uphill, through a barbed-wire fence, and into the forest about 100 yards. Leopards are reported to perform similar feats of strength, carrying heavy prey high into trees in order to eat undisturbed.
The coat of a jaguar is marked with black rosettes over an orangeyellow or tan background. Some jaguars have smaller spots not in the form of rosettes, and there is also a melanistic (black) version of the coat. The black jaguars (incorrectly called "black panthers") can appear in the same litter as spotted ones. However, a close look at their coats, particularly under an oblique light, reveals that the spots are still there, although hidden or masked by the dark background.
Adult jaguars can measure between 1.10 and 1.80 m from the head to the
base of the tail, which itself can measure between 440 and 560 cm. The longest jaguar on record measured 2.7 m (including the tail). Height at the shoulder has been measured between 68 and 75 cm. Jaguars can weigh up to 158 kg, with males weighing more than females. Central American jaguars tend to be smaller and lighter than their South American counterparts, there being considerable variation in the size and weight of jaguars from Central and South America.
Jaguars have large, strong canines and massive head musculature, which makes their faces appear very round. Their eyes shine a bright greenish-yellow under a light, and their ears are small and rounded.
I3/3, C1/1, P3/2, M1/1, for a total of 30 teeth.
Habitat and Distribution
Jaguars inhabit a wide range of habitats, from rain forests to wet grasslands, dry scrub lands, and even beaches. They are normally associated with bodies of water, such as rivers, streams, and lakes. Their large size requires abundant prey and large, relatively undisturbed territories in order to maintain viable populations. While jaguars will use the same habitats as pumas, pumas are usually displaced by jaguars when they occur in the same general area. They have been recorded from sea level to up to 3,800 meters in elevation, although they are usually found at less than 1,200 meters. Their range in Central America has been substantially reduced, with less than 30% of the original range still inhabited. They have become extirpated in the southwestern United States (although there has been at least one recent sighting in Arizona) and from Uruguay, so that now they are found from Mexico to Argentina.
Jaguars are very adaptable and exhibit diurnal and nocturnal habits throughout their range. They are solitary creatures that get together only during mating season. They keep territories which they patrol and mark with urine. They like to walk on man-made trails and are curious by nature, especially where they have not been hunted for a long time. As mentioned above, one female jaguar and her two cubs strolled into the Santa Rosa National Park camping area in the middle of the day, scattering a group of Boy Scouts and other visitors who took refuge in the park's administration building. Other jaguars have been seen walking along the beach at the same park during the early evening.
Jaguars are reported to like to rest on logs that overhang rivers, especially during the early morning hours. Karl Weidmann, a German naturalist, photographer, and filmmaker, reported (and recorded on film and in pictures) a close encounter with such a jaguar in Venezuela. It is documented in his excellent book Fauna de Venezuela (Weidmann, 1987). Normally shy due to overhunting, they are known to attack cattle and other livestock when desperate, usually in areas where their natural habitat has been reduced. In southern Nicaragua, for example, the combination of rapid colonization by farmers and excessive hunting of the jaguar's prey species (deer, tapirs, peccaries, agoutis, and others) has placed the remaining jaguars in a very precarious position. Seeking food, jaguars wander near settlements or farms, where they are quickly hunted down and killed by people using dogs and semiautomatic weapons.
Jaguars can roar, producing deep hoarse grunts that carry for hundreds of meters. Native jaguar hunters often imitate the call of the jaguar using a gourd or small drum over which a taut piece of rawhide has been fastened. A hole in the center, through which a string of leather or other material is passed and knotted, produces an uncannily similar series of grunts and calls which often elicit the curiosity of nearby animals. Alan Rabinowitz describes one of these callers and its use in his book Jaguar (Rabinowitz, 1986).
Jaguars are territorial, with the territories of males and females overlapping. A few studies have shown individual territories to be large, sometimes up to 25 square kilometers, with male territories about twice as large as those of females. It is not difficult to imagine, then, that the combination of large territorial needs and the continuing shrinkage of the jaguar's natural habitats place them in a no-win situation.
Jaguars are considered opportunistic carnivores, eating literally whatever is most abundant and easiest to catch. Besides cattle and horses, which are recent additions to the jaguar's diet, they normally feed on tapirs (Tapirus spp.), deer (both white-tailed and brocket deer), agoutis (Dasyprocta punctata), peccaries (Tayassu tajacu), arboreal mammals, bats, birds, fish, turtles, armadillos, anteaters, sloths, iguanas, coatis, snakes, caimans, and even fresh-water dolphins.
Jaguars are reported to breed year-round. During mating, they copulate frequently, as much as 100 times per day, although these copulations are very short (around nine seconds each). The offspring (one or two cubs, rarely three or even four) remain with the females until they are between 18 months and two years of age. They mature at about three years of age. Gestation lasts between 90 and 111 days, and the cubs weigh between 700 and 900 grams at birth. Cubs open their eyes after 13 days and keep the characteristic juvenile blue color of the irises almost until maturity. They den in caves, under fallen trees, in thickets, at riverbanks, under large rocks, or in other sheltered spaces. Adult females normally breed every two years.
Jaguars can live up to 11 years in nature, producing anywhere between four and eight cubs during their life span. This low rate of reproduction adds another element of risk to populations of jaguars in Central America. In captivity, jaguars have been known to live up to 23 years.
Central American jaguars have several threats, and their situation is grim in most of the region, except, perhaps, in Belize. Besides the drastic reduction of their natural range, their large territorial needs, the diminishing numbers of natural prey species, and the continuing pressure of the skin and trophy market make their plight in the region extremely serious. Habitat destruction due to the advance of agricultural frontiers is reported as the main cause of the decline of jaguar (and other feline) populations. The agricultural development carries with it other threats, such as overhunting of prey species, as well as increased hunting pressure on the jaguars themselves for their skins or because of the damage they cause to livestock. Attacks on humans are extremely rare throughout their range and clearly these attacks should not be a justification for eliminating them from a given area. Most documented attacks are from threatened, wounded, or old individual jaguars, particularly those who have lost teeth or the ability to overpower larger and faster prey.
Another serious threat to Central American jaguars is often called "the genetic factor." It has been calculated that to maintain a genetically healthy population of any animal species, a minimum number of 500 individuals should be present in a given area (Caughley and Gunn, 1996; Soulé, 1987). Given the territorial requirements and the other characteristics of jaguars, it is very difficult for them to maintain such numbers. The fragmentation of natural habitats in Central America precludes the maintenance of viable populations of jaguars in the region. Sadly, it is our opinion that free-living jaguars are doomed to disappear from most of Central America in the next 50 years or less.
Jaguars are listed under CITES Appendix I and as endangered in all of the countries in Central America except Belize. (See "The Future of Central American Carnivores" for an explanation of CITES.)