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Excerpted from The Costa Rica Handbook, by Christopher Baker

Anyone who has traveled in the tropics in search of wildlife can tell you that disappointment comes easy, and often at considerable expense. But Costa Rica is one place that lives up to its word. You don't need to venture far to experience the nation's full panoply of magnificent wildlife. Costa Rica is nature's live theater where the actors aren't shy.

Crocodiles and Caimans

Many travelers visit Costa Rica in the hope of seeing crocodiles and caimans, modest-sized relatives. One of the smallest of western crocodilians--no more than two meters long--and possibly the most abundant in existence today, the speckled caiman is still relatively common in parts of wet lowland Costa Rica on both the Atlantic and Pacific coasts. Palo Verde and Tortuguero are both good places to spot them in small creeks, playas, and brackish mangrove swamps, or basking on the banks of streams and ponds.

The scales of the caiman take on the blue-green color of the water it slithers through. Such camouflage and even the ability to breathe underwater, through raised nostrils, have not protected the caiman. Their nests are heavily disturbed by dogs, foxes, tegu lizards, and humans. And increasingly they are being sought for their skins, which are turned into trivia. Ironically, this is easing the pressure on the crocodiles, which are fast disappearing as humanity takes their hides and habitats.

The crocodile exists in precariously low numbers along both coasts, and the only healthy population is in Corcovado National Park, in the Pacific southwest. Only three species of crocodiles -- the saltwater croc of Southeast Asia, Africa's Nile crocodile, and the American alligator -- are considered man-killers, but it's still a wise idea to check with locals or park rangers before swimming in coastal estuaries and lagoons.

For all their beastly behavior, crocodiles are devoted parents. And despite being relics from the age of the dinosaurs, croc brains are far more complex than those of other reptiles. Perhaps because crocodiles have ugly toothy leers and a stigma of primeval wickedness, there isn't the same love of crocs that has brought international support for the turtles, and their future is much less secure. As biologist David Janzen says: "We may never again see the huge four-meter animals that used to terrify the campesinos and eat their dogs."
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No visit to Costa Rica would be complete without seeing any of its four species of monkeys: the cebus (or capuchin), howler, spider, and squirrel. Along with approximately 50 other species, they belong to a group called New World monkeys, which evolved from a single simian group that appeared about 40 million years ago in Africa and Asia. Some of these early primates migrated to North America and then down the land bridge to Central and South America.

Though the North American monkeys gradually died out, their southern cousins flourished and evolved along lines that differ markedly from those of their ancestors in the Old World. While African and Asian monkeys have narrow noses with nostrils that point down (much like human noses), New World monkeys evolved broad, widely spaced nostrils. New World females, too, evolved a singular ability to bear twins. And, perhaps most importantly, some New World species--notably the cebus, howler, and spider monkeys--developed long prehensile tails for added purchase and balance in the high treetops.

They inhabit a wide range of habitats, from the rainforest canopy to the scrubby undergrowth of the dry forests, though each species occupies its own niche and the species seldom meet. Together, they are the liveliest and most vocal jungle tenants. Beyond the reach of most predators, they have little inhibition about announcing their presence with their roughhousing and howls, chatterings, and screeches. The sudden explosive roar of the howler monkey -- a sound guaranteed to make your hair stand on end -- is said to be the loudest sound in the animal kingdom.

The distinctive-looking capuchin is the smartest and most inquisitive of Central American simians. It derives its name from its black body and monklike white cowl. If they look familiar to you, you've probably seen them dancing at the end of a tether at street fairs in Europe or South America--they're the little guys favored by organ-grinders worldwide. Capuchins range widely throughout the wet lowland forests of the Caribbean coast and the deciduous dry forests of the Pacific northwest below 1,500 meters. Two excellent places to see them are Santa Rosa and Manuel Antonio national parks, where family troops are constantly on the prowl, foraging widely through the treetops and over the forest floor.

These opportunistic feeders are fun to watch as they search under logs and leaves or tear off bark as they seek out insects and small lizards soon after dawn and again in late afternoon. Capuchins also steal birds' eggs and nestlings. Some crafty coastal residents, not content with grubs and insect larvae, have developed a taste for oysters and other mollusks which they break open on rocks. The frugal capuchin sometimes hoards his food for "rainy days." While their taste is eclectic, they are fussy eaters: they'll meticulously pick out grubs from fruit, which they test for ripeness by smelling and squeezing. And capuchins are not averse to crop-raiding, especially corn, as the farmers of Guanacaste will attest.
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There's something endearing about the leaf-cutting ant (Atta cephalotes), a mushroom-farming insect found in lowland forests throughout Costa Rica, carrying upright in its jaws a circular green shard scissored from the leaves of a plant. At some stage in your travels you're bound to come across an endless troop of "media" workers hauling their cargo along jungle pathways as immaculately cleaned of debris as any swept doorstep.

The nests are built below ground, sometimes extending over an area of 200 square meters, with galleries to a depth of six meters. Large nests provide a home for up to five million insects. (All ant societies are composed entirely of females; males exist only to fertilize the queen and then die. And only the queen, who may boast a thousand times the body weight of a minor worker, is fertile. Hence, all other ants in the colony are her daughters.) They set off from their nests, day and night, in long columns to demolish trees, removing every shoot, leaf, and stem section by tiny section and transporting them back to their underground chambers.

They don't eat this material. Instead, they chew it up to form a compost on which they cultivate a nutritional breadlike fungus whose tiny white fruiting bodies provide them with food. The species has evolved different physical castes, each specializing in its own social tasks. Most of the workers are tiny minors ("minimas"), which tend the nest. The cutting and carrying is performed by intermediate-size workers ("medias"), guarded by ferocious-looking "majors" about three-quarters of an inch long.

The most terrifying ants of all are the army ants, which march through the forest with the sole intent of turning small creatures into skeletons in a few minutes. They produce a faint hissing sound and distinct ant-army odor. They're like a wolf pack, but with tens of thousands of miniature beasts of prey which merge and unite to form one great living creature.

Hollywood images of mammals and even humans fleeing madly before them are mostly imagination run wild. In truth, while the ants advance across the forest floor driving small creatures in front of them, humans and other large creatures can simply step aside and watch the column pass by--this can take several hours. Even when the ants raid human habitations, people can simply clear out with their foodstock while the ants clean out the cockroaches and other vermin as thoroughly as any exterminator might.

The army ants' jaws are so powerful that Indians once used them to suture wounds: the tenacious insect was held over a wound and its body squeezed so that its jaws instinctively shut, clamping the flesh together. The body was then pinched off.

Larvae carried by workers produce pheromones which stimulate the army to keep on the move. When the larvae begin to pupate and no longer exude their chemical messages, the ants bivouac in a vast ball in a hollow. They actually cling to one another and make a nest of their bodies, complete with passageways and chambers where the eggs are deposited. Once the queen lays her eggs and these hatch as larvae, a new generation of workers and soldiers synchronistically emerges from the stored pupae. The larvae begin to secrete their characteristic pheromone, and the army is again stimulated to march off and terrorize the bush.
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Poison Dart Frogs

Of all Central America's exotic species none are more colorful -- literally and figuratively -- than the "poison-arrow" frogs, of the type from which Indians extract deadly poisons with which to tip their arrows. Frogs are tasty little fellows to carnivorous amphibians, reptiles, and birds. Hence, in many species, the mucous glands common in all amphibians have evolved to produce a bitter-tasting poison.

In Central and South America at least 20 kinds of frogs have developed this defense still further: their alkaloid poisons are so toxic that they can paralyze a large bird or small monkey immediately. Several species -- the dendrobatids, or poison-arrow frogs, which are confined to Costa Rica -- produce among the most potent toxins known: atelopidtoxin, bufogenin, bufotenidine, and bufotoxin. Pity the poor snake that gobbles up Dendrobatis granuliferus, a tiny, bright green, red, and black frog which inhabits the lowland forests of the Golfo Dulce region (it is commonly seen on forest floors of Corcovado National Park). Another species, Bufo marinus, can even squirt its poison in a fine spray. And some species' eggs and tadpoles even produce toxins, making them unpalatable, like bad caviar!

Of course, it's no value to an individual frog if its attacker dies after devouring the victim. Hence, many have developed conspicuous, striking colors--bright yellow, scarlet, purple, and blue, the colors of poison recognized throughout the animal world--and sometimes "flash colors" (concealed when at rest but flashed at appropriate times to startle predators) that announce, "Beware!" These confident critters don't act like other frogs either. They're active by day not night, moving boldly around the forest floor, "confident and secure," says one writer, "in their brilliant livery."

Perhaps the most famous poison frog species is the rare golden toad, found only in the Monteverde Cloud Forest Reserve. In fact, the montane rainforest reserve owes its existence in part to the discovery of Bufo periglenes. This brilliant, neon orange arboreal toad--discovered in 1964 and so stunning that one biologist harbored "a suspicion that someone had dipped the examples in enamel paint"--is not easily seen, despite being one of the most brightly colored animals in the world. The golden toad is so seldom seen that some naturalists speculate that it may now exist only on the cover of tourist brochures.

April through May, golden toads go looking for love in the rainpools of scarlet bromeliads that festoon the high branches. They're easily distinguished: the males are the orange ones; females are yellow and black with patches of scarlet. Here, high in the trees, tadpoles of arboreal frogs wriggle about.

Few Costa Rican frog species breed in permanent bodies of water, where fish predation is intense. Above 1,500 meters, where there are no native fish species, stream breeding is more common (although the introduction in recent years of trout into upland streams already threatens whole frog populations).

The frogs instead have evolved away from dependence on bodies of permanent water. Many species, particularly the 39 species of hylids, spend their entire lives in the tree canopies where they breed in holes and bromeliads. (The hylas have enlarged suction-cup pads on their toes. They often catch their prey in midair leaps: the suction discs guarantee surefooted landings.) Others deposit their eggs on vegetation over streams; the tadpoles fall when hatched. Others construct frothy foam nests which they float on pools, dutifully guarded by the watchful male.

Some rainforest species, such as the diminutive and warty eleutherodactylus -- its name is longer than its body -- live on the ground, where they lay their eggs in moist cups of leaves. The tadpole develops fully within the egg sac before emerging as a perfect, if tiny, replica of its parents: it and 12 of its siblings could easily fit on a man's fingernail. Some tadpole species -- the Hyla zeteki, for example -- are carnivorous: they eat other frogs' tadpoles. Others, like the smoky frog (Leptodactylus pentadactylus) are even cannibalistic! This aggressive giant (adults can grow up to eight inches long) can eat snakes up to 20 inches long. It, too, can emit a poisonous toxin, to which some snakes are immune. If the smoky frog's loud hissing, inflated body, and poisonous secretions don't manage to scare off its predator, it has another ingenious defense: when captured it emits a loud scream.
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The quetzal, or resplendent trogon, is a rare jewel of the bird world. Many birdwatchers travel to Costa Rica simply to catch site of this magnificent bird. What this pigeon-sized bird lacks in physical stature it makes up for in audacious plumage: vivid, shimmering green which ignites in the sunshine, flashing emerald to golden and back to iridescent green. In common with other bird species, the male outshines the female. He sports a fuzzy pink punk hairdo, a scintillating crimson belly, and two brilliant green tail plumes up to 24 inches long, edged in snowy white and sinuous as feather boas.

Its beauty was so fabled and the bird so elusive and shy that early European naturalists believed the quetzal was a fabrication of Central American natives. In 1861, an English naturalist, Osbert Salvin, wrote that he was "determined, rain or no rain, to be off to the mountain forests in search of quetzals, to see and shoot which has been a daydream for me ever since I set foot in Central America." Salvin, the first European to record observing a quetzal, pronounced it "unequaled for splendour among the birds of the New World," and promptly shot it. During the course of the next three decades, thousands of quetzal plumes crossed the Atlantic to fill the specimen cabinets of European collectors and adorn the fashionable milliners' shops of Paris, Amsterdam, and London. Salvin redeemed himself by authoring the awesome 40-volume tome Biologia Centrali Americana, which provided virtually a complete catalog of neotropical species.

The quetzal has long been revered in Guatemala, where the bird graces the national shield, flag, postage stamps, and currency (which happens to be called the quetzal). It is pleasing to know that the former center of the Mayan empire still honors the magnificent bird. Early Mayans and Aztecs worshiped a god called Quetzalcoatl, the Plumed Serpent, and depicted him with a headdress of quetzal feathers. The bird's name is derived from quetzalli, an Aztec word meaning "precious" or "beautiful."

Mayans considered the male's iridescent green tail feathers worth more than gold, and killing the sacred bird was a capital crime. Quetzal plumes and jade, which were traded throughout Mesoamerica, were the Mayans' most precious objects. It was the color that was significant: "Green--the color of water, the lifegiving fluid. Green, the color of the maize crop, had special significance to the people of Mesoamerica," says Adrian Digby in his monograph Mayan Jades, "and both jade and the feathers of the quetzal were green."

During the colonial period, the indigenous people of Central America came to see the quetzal as a symbol of independence and freedom. Popular folklore relates how the quetzal got its dazzling blood-red breast: in 1524, when the Spanish conquistador Pedro de Alvarado defeated the Mayan chieftain Tecun Uman, a gilt-and-green quetzal alighted on the Indian's chest at the moment he fell mortally wounded; when the bird took off again, his breast was stained with the brilliant crimson blood of the Mayan.

Archaeologists believe that the wearing of quetzal plumes was proscribed, under pain of death, for use by Mayan priests and nobility: it became a symbol of authority vested in a theocratic elite, much as only Roman nobility were allowed to wear purple silks.

Although Costa Ricans don't worship the quetzal with the same fervor as pre-Columbian Guatemalans, the bird is most easily seen in Costa Rica, where it is protected in four national parks -- Braulio Carrillo, Poás, Chirripó, La Amistad -- and the Monteverde and Los Angeles cloud forest reserves. Everywhere throughout its 1,000-mile range (from southern Mexico to western Panama) it is endangered due to loss of its cloud-forest habitat. This is particularly true of the lower forests around 1,500 to 2,000 meters to which families of quetzals descend during breeding season (March-June), and where they seek dead and decaying trees in which to hollow out their nests. This is the best time to see narcissistic males showing off their tail plumes in undulating flight, or launching spiraling skyward flights which presage a plummeting dive with their tail feathers rippling behind, all part of the courtship ritual.

At other times, the wary birds aren't easily spotted. Their plumage offers excellent camouflage under the rainy forest canopy. They also sit motionless for long periods, with their vibrant red chests turned away from any suspected danger. If a quetzal knows you're close by and feels threatened, you may hear a harsh weec-weec warning call and see the male's flicking tail feathers betray his presence. The quetzal's territory spans a radius of approximately 300 meters, which the male proclaims each dawn through midmorning and again at dusk with a telltale melodious whistle -- a hollow, high-pitched call of two notes, one ascending steeply, the other descending -- repeated every eight to 10 minutes.

Nest holes (often hollowed out by woodpeckers) are generally about 30 feet from the ground. Within, the female generally lays two light-blue eggs, which take about 18 days to hatch. Both sexes share parental duties. By day, the male incubates the eggs while his two-foot-long tail feathers hang out of the nest. At night, the female takes over.

Although the quetzal eats insects, small frogs, and lizards, it enjoys a penchant for the fruit of the broad-leafed aguacatillo (a kind of miniature avocado in the laurel family), which depends on the bird to distribute seeds. The movement of quetzals follows the seasonal fruiting of different laurel species. Time your birdwatching visit, if possible, to coincide with the quetzals' rather meticulous feeding hours, which you can almost set your watch by. They're fascinating to watch feeding: an upward swoop for fruit is the bird's aerial signature.
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Although rarely seen by the casual tourist in Costa Rica, snakes make up almost half of all reptile species in the nation (135 species, 17 poisonous). It is a fortunate traveler indeed who gets to see in the wild the fantastically elongated, I-beam-shaped chunk-headed snake, with its catlike elliptical eyes; the slender, beak-nosed bright green vinelike vine snake; or the relatively benign boa constrictor. Many neotropical snake species inhabit a wide range of Costa Rican environments, and wherever you are in the country there are sure to be snakes about. Not that you should worry. Fewer than 500 snakebites are reported each year, and less than three percent of these are fatal. Most bites occur among farmworkers.

Still, caution is always the watchword. Never reach into holes or under rocks, debris, or forest-floor leaf litter without first checking with a stick to see what might be quietly slumbering there. And remember that many snakes are well-camouflaged arboreal creatures, which snooze on branches, so never reach for a branch without looking. You should even be cautious when peering inside bromeliads: the dark-colored chunk-headed snake likes to doze inside the moisture-collecting plants during the dry season.

Among the more common snake species you are likely to see are the wide-ranging boas which, with luck, you might spot crawling across a cultivated field or waiting patiently in the bough of a tree in wet or dry tropical forest, savannah, or dry thorn scrub. Wild boas vary in temperament and some are aggressive. Though not poisonous, they are quite capable of inflicting serious damage with their large teeth and will not hesitate to bite. Heaven forbid a full-grown adult (three meters or more) should sink its teeth in sufficiently to get its constricting coils around you!

A species of tropical rattlesnake is also found in Guanacaste and a few relic areas of the Meseta Central. It produces a venom considerably more toxic than its North American cousin -- blindness and suffocation are typical effects on humans -- and it rarely uses its well-developed rattle to warn off the unwary.

Among the more colorful snakes are the four species of coral snakes, with small heads, blunt tails, and bright bands of red, black, and yellow or white. These highly venomous snakes (often fatal to humans) exhibit a spectacular defensive display when approached: they flatten their bodies and snap back and forth while alternately hiding then swinging their heads side to side and coiling and waving their tails.

Along the Pacific beaches, you may sometimes encounter venomous pelagic sea snakes, a yellow-bellied and black-backed serpent closely related to terrestrial cobras and coral snakes. This gregarious snake has developed an oarlike tail to paddle its way through the ocean. It tends to drift passively with its buddies among drift-lines of flotsam, where it feeds on small fish.

The most talked-about snake in Central America is the fer-de-lance, much feared for its aggressiveness and lethal venom. One of several Central American pit vipers -- another is the bushmaster -- the fer-de-lance can grow to a length of three meters and is abundant throughout the country, particularly in overgrown fields and rivercourses in drier lowland regions. Costa Ricans call this lethal creature terciopelo, Spanish for "velvet." As juveniles, fer-de-lance are arboreal critters that feed on lizards and frogs, which they attract with a yellow-tipped tail. As adults, they come down to earth, where they move about at night and, by daylight, rest in loose coils of burnished brown on the forest floor.

Give the fer-de-lance a wide berth! Unlike other vipers, the fer-de-lance will bite with little provocation. The snake's powerful venom dissolves nerve tissue and destroys blood cells and artery walls; those fortunate enough to survive may suffer paralysis or tissue damage so massive as to require amputation of the bitten limb.

The Serpentario in San José is a good place to learn to identify snakes and their habits and habitats. There is also a snake laboratory at the Clodomiro Picando Institute in Coronado, where you can watch snakes being milked for venom.
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With nearly 1,000 identified species (approximately 10 percent of the world total), Costa Rica is a lepidopterist's paradise. You can barely stand still for one minute without checking off a dozen dazzling species: metallic gold riondinidae; delicate black-winged heliconius splashed with bright red and yellow; orange-striped paracaidas; and the deep neon-blue flash of morphos fluttering and diving in a ballet of subaqueous color. The marvelously intricate wing patterns are statements of identity, so that individuals may recognize those with whom mating may be fertile.

Not all this elaboration has a solely sexual connotation. Some butterflies are ornately colored to keep predators at bay. The bright white stripes against black on the zebra butterfly (like other members of the Heliconid family), for example, tell birds that the butterfly tastes acrid. There are even perfectly tasty butterfly species, which mimic the Heliconid's colors, tricking predators to disdain them. Others use their colors as camouflage so that at rest they blend in with the green or brown leaves or look like the scaly bark of a tree. Among the most intriguing, however, are the owl-eye butterflies, with their five-inch wingspans and startling eye spots. The dull, blue-gray Caligo memnon, the cream owl butterfly, is the most spectacular of the owl-eyes: the underside of its wings is mottled to look like feathers and boasts two large yellow-and-black "eyes" on the hind wing, which it displays when disturbed.

The best time to see butterflies is in the morning, when most species are active. A few are active at dawn and dusk, and there is even one species that is active by night. In general, butterfly populations are most dense in June and July, corresponding with the onset of the rainy season on the Pacific side. Butterfly migrations are also common. Like birds, higher-elevation species migrate up and down the mountains with changes in local weather. The most amazing migration -- unsurpassed by any other insect in the Neotropics -- is that of the kitelike uranidae (this black and iridescent green species is actually a moth that mimics the swallowtail butterfly), in which millions of individuals pass through Costa Rica heading south from Honduras to Colombia.
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About the author

Christopher Baker was born and raised in Yorkshire, England. After receiving a B.A. in Geography at University College London (including two Sahara research expeditions and an exchange program at Krakow University, Poland), he earned Masters degrees in Latin American Studies from Liverpool University, and in Education from the Institute of Education, London University. Baker began his writing career in 1978 as a Contributing Editor on Latin America for Land and Liberty, a London-based political journal. In 1980, he received a Scripps-Howard Foundation Scholarship in Journalism to attend the University of California, Berkeley. Since 198, he has made his living as a professional travel and natural-sciences writer, and has been published in over 150 publications worldwide. Baker is a member of the Society of American Travel Writers and has received several awards for outstanding writing, including the Lowell Thomas Travel Journalism award. He lives in Oakland, California.

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