"If I discover within myself a desire which no experience of this world can satisfy,

the most probable explanation is that I was made for another world."

    Clive Staples Lewis (1898-1963)

    Anglican philosopher and novelist

 

Machaonides swallowtail, Heraclides machaonides. Bonao, central Dominican Republic, Hispaniola.

 


 

The Swarming Hordes

 

CLASS INSECTA: INSECTS

 

    When I was a boy, biologists had calculated that the grand total of organic species - plant and animal - on Earth was about one million. More recent studies have given us a better understanding of the actual number of life-forms on our planet. Especially since new techniques have allowed ecologists and taxonomists to have a better access to one of the last biological frontiers, the rain forest's canopy, it is now believed that the number of living species of insects alone is between four and 30 million. Although these estimates are far from certain, and even assuming only four million species of insects, add to that the number of parasites - bacterial, protozoan, fungal, nematodan, acarine, and many of which are highly host-specific - that plague those same insects, and then the hyperparasites that the infest the parasites, and that will give us an ever better glimpse of the staggering biological diversity on Earth. (In fact, a very large percentage of the species on the planet are parasites of other organisms).

 

    In any case, and simply put, there are more species of insects on this planet than all others combined.

 

    Insects are considered a class (Insecta) within the subphylum Hexapoda ("six-legged"), together with the classes Diplura, Protura, and Collembolla. Insects are usually the most abundant macroscopic animals in all terrestrial ecosystems. They are also present, whether as eggs, larvae, or adults, in practically every freshwater environment though, interestingly, only a very few species are marine, and even those live only in coastal areas and are never pelagic (none lives in the open ocean). None are totally aquatic throughout their lives, since all adult insects need to breathe gaseous oxygen, even those that do spend their adult lives in water.

 

Larva of Puerto Rican rhinoceros beetle, Strategus quadrifoveatus, showing on its sides the spiracles leading to its tracheae. All insects need to breathe gaseous oxygen, at least as adults.

Fort Buchanan, Guaynabo, north-eastern Puerto Rico.

 

    In turn, Insecta is divided into three subclasses. Two of these, Archaeognatha (bristletails) and Thysanura (silverfish) are the most primitive of all insects, and are seldom-seen denizens of damp, dark places. The remaining subclass, Pterygota, contains all the other species. These are the "winged insects", including forms that have secondarily lost their wings and capacity for flight.

 

    Although the relationships of insects are obscure, they seem to be most closely related to crustaceans than to any other group of living arthropods. Perhaps arising during the Devonian Period, they really exploded in terms of numbers of individuals and species later, coming into their own during the Carboniferous.

 

    Some insects rely on color to advertise themselves to others of their kind or to other types of animals. Also, aposematic combinations of black, red, orange, and yellow are widely recognized in the animal kingdom as a warning meaning "you better not eat me/not bother me - I taste awful/I am dangerous." (The blacks, reds, and yellows of many wasps and bees are well-known examples of this phenomenon).

 

    Other insects depend on camouflage to blend with their surroundings, imitating the colors and shapes of the substrate on which they live. Some have carried this to such extremes that they are practically indistinguishable from twigs, lichens, or even flowers.

 

    Alone among invertebrates, insects developed the capacity for true flight and where the first organisms in the history of Earth to take to the air, possibly in the early Carboniferous Period. Some orders, like the primitive Thysanura (silverfish) have no wings, while most insects have two pairs of wings. In the case of dipterans (flies and allies) only two wings are functional for flight, while the other two has been modified as instruments for balance. Certain groups, like some walking sticks and the worker and soldiers classes of ant and termite colonies, have secondarily lost their wings altogether, having evolved from winged ancestors. A few extinct groups of insects had three pairs of functional wings, but this trait was lost early in favor of only four wings.

 

    The wings of insects evolved from appendages that possibly had at first the appearance of legs. Unlike the wings of birds, bats, and pterosaurs, those of insects have no muscle and are mostly rigid. The animal flies by beating and rotating them with the use of muscles that alter the shaped of the bodily wall, thus bringing movement to the wings.

 

    Some small insects, like midges and mosquitoes, can beat their wings tens of thousands of times per second. Many larger ones, like butterflies and dragonflies, beat them at a comparatively much more leisurely rate, and can glide of them when unhurried.

 

Order Blattodea: Cockroaches

 

    Cockroaches are usually associated with dirt and germs, but in reality most species live far away from human habitations, in forests, grasslands, and even caverns. Many species are omnivorous, eating whatever they can find (which is part of the reason why a number of species feel so comfortable in your kitchen and pantry).

 

    In spite of great morphological differences among them, this order of insects is actually closely related to three others: crickets and grasshoppers, mantids, and walking sticks. More distantly, they are also kin to the termites.

 

Puerto Rican cave cockroach, Aspiduchus cavernicola. Cueva de la 22, Cabo Rojo, south-western Puerto Rico.

This species lives exclusively in caves and caverns, where they feed on dead bats and other carrion.

 

Arboreal cockroach, Panchlora sp. El Yunque National Forest, north-eastern Puerto Rico.

 


Mating pair of cockroaches, probably Epilampra wheeleri. Carite State Forest, east-central Puerto Rico.
The color and shape of this species blend with the forest's leaf litter in with it lives.

 

Arboreal cockroach, Pelmatosilpha coriacea. El Yunque National Forest, north-eastern Puerto Rico.

 

Arboreal cockroach, species undetermined. Grand Etang National Park, central Grenada, Lesser Antilles.

 

Arboreal cockroach, probably Pelmatosilpha sp. Kingstown Botanical Garden, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.

 

Cockroach, species undetermined. Windsor, north-central Jamaica.

 

Cockroach, species undetermined. Mount Sage, west-central Tortola, British Virgin Islands.

 

Order Coleoptera: Beetles

 

    Beetles comprise the largest group of insects, indeed the single largest order of things living. They can be herbivores, predators, or carrion eaters, and comprise an important role in the terrestrial ecosystems of the world, for their great abundance and variety of species.

 

    Their first pair of wings is highly chitinous and are no really used from flight. The second pair, which during rest is folded under the first, is the one used for that purpose.

 

Cerambycid beetle, species undetermined. Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Cerambycid beetle, Stenodontes maxillosus. Saint Bartholomew, Lesser Antilles.

(Photograph courtesy of Mr. Karl Questel).

 

Cerambycid beetle, probably Stenodontes sp. Susua State Forest, south-western Puerto Rico.

 

Cerambycid beetle, Solenoptera thomae, female. Susua State Forest, south-western Puerto Rico.

 

Cerambycid beetle, Lagochirus araneiformis. Barrenspot, central Saint Croix, United States Virgin Islands.

 

Cerambycid beetle, Plinthocoelium domingoensis. Del Este National Park, eastern Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Piero Fariselli).

 

Cerambycid beetle, Eburia quadrimaculata. Guana Island, British Virgin Islands.

 

Cerambycid beetle, species undetermined, resting on a stem. San Juan Botanical Garden, San Juan, north-eastern Puerto Rico.

 

Firefly, probably Photinus sp. Toro Negro State Forest, central Puerto Rico.

 

Firefly, Photuris jamaicensis. Windsor, north-central Jamaica.

 

Some elaterids have bioluminescent organs, frequently paired and on top of the thorax, as can be seen in this individual.

These emit a green light at night or when the animal feels threatened.

Members of this group are frequently called "click beetles" because they can cock and then suddenly release a hinge between

their thoraxes and abdomens. An individual placed on its back can thus jump in the air several inches, until it falls on its legs, once a gain.

This is Pyrophorus luminosus. Carite State Forest, east-central Puerto Rico.

 

Chrysomelid beetles, probably Altica occidentalis, feeding on leaves.

Guanica State Forest, south-western Puerto Rico.

Members of this and related genera often are tiny creatures, thus frequently called flea beetles.

 

Chrysomelid beetles, probably Altica occidentalis, feeding on leaves.

El Yunque National Forest, north-eastern Puerto Rico.

 

Tortoise beetles (family Chrysomelidae), possibly Charidotella sp., male (dull orange and black) and females (iridescent purple and greenish yellow).

Camp Santiago, Salinas, south-eastern Puerto Rico.

Chrysomelids are frequently called "tortoise beetles" for their general shapes.

 

The overhanging shape of the thorax and elytra of Charidotella sp. allow them

to fit closely against their substrate as protection against attackers like ants and similar insects.

Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Scarabeid beetle, species undetermined. Eggleston, south-central Dominica, Lesser Antilles.

 

Scarabeid beetle, species undetermined. Guilarte State Forest, central Puerto Rico.

 

Puerto Rican rhinoceros beetles, Strategus quadrifoveatus, larva, male (two center photographs), and female. Fort Buchanan, Guaynabo, north-eastern Puerto Rico.

 

Puerto Rican May beetle, Phyllophaga portoricensis. El Yunque National Forest, north-eastern Puerto Rico.

 

May beetle, Phyllophaga sp. Carolina, north-eastern Puerto Rico.

 

Tiger beetles (family Cicindelidae), Cicindela boops. Guanica State Forest, south-western Puerto Rico.

 

Carabid beetle, Galerita sp. beetle. Negril, western Jamaica.

 

Tenebrionid beetle larva, species undetermined. Guilarte State Forest, west-central Puerto Rico.

 

Tenebrionid beetles, probably Tenebrio sp. Barrenspot, Central Saint Croix, United States Virgin Islands.

 

Weevil, Diaprepes abbreviatus. Charlotte Amalie, southern Saint Thomas, United States Virgin Islands.

This animal is found in the insular banks of Hispaniola, Puerto Rico, and some of the Lesser Antilles.

More than 65000 species of weevils are known, and surely many more await discovery, possibly a quarter of a million.

They are the largest family (Curculionidae) of the largest order (Coleoptera) of the largest class (Insecta)

of the largest superclass (Hexapoda) of the largest phylum (Arthropoda) of the largest kingdom (Metazoa) of biological organisms.

 

Another morph of Diaprepes abbreviatus. El Yunque National Forest, north-eastern Puerto Rico.

As with some other beetles, and in a manner similar to butterflies, the color patters of these weevils is derived

from a thousands of microscopic scales that cover their elytra. The dark lines are actually places where the scales

fall off, and show the underlying color of the animal. This species is a pest of several crops, and has caused great

economic losses in places like Florida, United States, where it has been introduced.

 

A weevil, Diaprepes maugei, hitches a ride atop a walking stick, Lamponius sp. Toro Negro State Forest, central Puerto Rico.

 

Weevil, possibly Diaprepes glaucus. Toro Negro State Forest, central Puerto Rico.

 

A Diaprepes weevil spreads its wings upon take off. Port Antonio, north-eastern Jamaica.

 

Weevil, Exophthalmus quindecimpunctatus. Cambalache State Forest, north-central Puerto Rico.

(Photograph courtesy of Dr. Luis O. Nieves).

 

Weevil, probably Polydrusus sp. Rio Grande, north-eastern Puerto Rico.



Weevil, species undetermibed. Soufriere Botanical Gardens, south-eastern Saint Lucia, Lesser Antilles.


Twig-mimicking weevil, possibly Brentus sp. Eggleston, south-central Dominica, Lesser Antilles.

 

Order Dermaptera: Earwigs

 

    These peculiar insects, often called earwigs, have a pair of forceps on the end of their abdomens, somewhat like a crab's pincers. They feed on dead vegetable matter and carrion, and some species can be quite large for a member of their class. The forceps are used in copulation and as a defensive apparatus, capable of inflicting surprising but otherwise harmless pinch.

 

Earwig, possibly Euborellia caraibea. Guilarte State Forest, west-central Puerto Rico.

 

Order Diptera: Flies, Mosquitoes, and Their Kin

 

    Flies and their kin have only two functional wings. The second pair has been modified as balancing organs used to keep the right orientation during flight.

 

    Although flies are not usually considered to be predators, some are, indeed. Asilids (widely known as "robber flies") for example, have evolved to prey on other insects. Their proboscii (basically the same structures as those of blood-sucking mosquitoes) are inserted into the bodies of their hapless victims in order to suck their fluids. Large individuals can inflict a nasty prick on a human finger, as well.

 

    Most dipterans, however, feed on vegetable matter or carrion, while mosquitoes are well-known for being parasites of vertebrates. Some of the latter are vectors of important, and sometimes deadly, diseases like malaria, and hemorrhagic fevers like dengue.

 

Robber fly (family Asilidae), Proctacanthus sp., male. Enriquillo National Park, south-western Dominican Republic, Hispaniola.

 

Robber flies, Efferia sp., male, and female. Guanica State Forest, south-western Puerto Rico.

 

Robber fly, Ommatius marginellus, male. Caja de Muerto Nature Reserve, off southern Puerto Rico.



Robber fly, female, species undetermined. Soufriere Botanical Gardens, souther-eastern Saint Lucia, Lesser Antilles.

 

Micropezid flies, Taeniaptera lasciva, females. Aguas Buenas, east-central Puerto Rico.

Members of this family often mimic wasps or ants. Their forelegs are usually shortest and possess white tips.

 

Mating pair of flower flies, Meromacrus cintus. El Yunque National Forest, north-eastern Puerto Rico.

These and other species of dipterans earn some protection from predators by mimicking wasps and bees.

 

Flower flies, Palpada vinetorum. Pinones State Forest, north-eastern Puerto Rico.

 

Carrion fly, Phaenicia sp., (family Calliphoridae), male feeding on nectar. Aguas Buenas, east-central Puerto Rico.

 

Deerfly (family Tabanidae, subfamily Crysopsinae), Chrysops variegata, feeding on my blood.

Camuy, north-western Puerto Rico.

 

Soldier fly (Family Stratiomyidae), possibly Hedridiscus sp. Loiza, north-eastern Puerto Rico.

 

Mosquito, species undetermined.

Camp Santiago, Salinas, south-eastern Puerto Rico.

 

An adult midge has just emerged from its pupa, its as yet transparent body not hard enough for flight.

Species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Crane fly (family Tipulidae), species undetermined.

El Yunque National Forest, north-eastern Puerto Rico.

 

Mating pair of crane flies (family Tipulidae), species undetermined.

El Yunque National Forest, north-eastern Puerto Rico.

 

Mosquitoes do not always feed on the blood of mammals and birds. They will also feed off reptiles,

like this one does as it sucks the blood of a sleeping juvenile female Puerto Rican giant anole, Anolis cuvieri.

Mata de Platano Field Station, Arecibo, northern Puerto Rico.

 

Some parasitic flies have developed bizarre lifestyles. The two "ticks" on the abdomen of this

tettigonid grasshopper are actually ceratopogonid midges, feeding on the hemolymph of their host.

Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

 

Some flies are endoparasites (they live inside the tissues of living hosts).

These are the maggots of the bot fly Philornis pici. (family Cuterebridae). They have developed under

the skin of some birds, in this case a fledging Puerto Rican woodpecker (Melanerpes portoricensis).

The bird has died after more than 15, two centimeter-long maggots have consumed much the muscle tissues of the live bird.

Once their host is dead, the larvae leave it to pupate on the ground.

 

Order Hemiptera: Bugs, Cicadas, Planthoppers, and Their Kin

 

    Bugs and their allies have piercing mouth parts used to suck fluids of their prey or pf plants. Some species are important vectors of both animal and plant diseases. Among these last are the reduviid bugs that are the vectors of the trypanosome protozoans that cause the dreaded Leishmaniasis, Chagas disease, and sleeping sickness. Others are serious agricultural pest, and a number of them are predators of other insects.

 

    While most species are terrestrial, members of several families are aquatic, yet retaining the capacity for flight if local conditions become inadequate for survival. Some of these are the giant water bugs, backswimmers, water skaters, and water "scorpions".

 

Pyrrhocorid bugs, probably Dysdercus sp. Crown Mountain, central Saint Thomas, United States Virgin Islands.

 

Pyrrhocorid bugs, Dysdercus sanguineus neglectus. Guanica State Forest, south-western Puerto Rico.

 

Pyrrhocorid bugs, Dysdercus andreae. Guanica State Forest, south-western Puerto Rico.

 

Scutellerid ("jewel") bug, Pachycoris fabricii. El Yunque National Forest, north-eastern Puerto Rico.

(Photograph courtesy of Dr. Javier Mercado).

 

Scutellerid bug, Pachycoris sp. Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Some bugs have variant morphs. This is the pentatomid Vulsirea nigrorubra.

First two photographs: El Yunque National Forest, north-eastern Puerto Rico.

Last photograph, Guanica State Forest, south-western Puerto Rico.

This species is probably distasteful or even toxic, as advertised by its bright colors.

 

Pentatomid bug, Loxa viridis. Guanica State Forest, south-western Puerto Rico.

 

Pentatomid bugs, species undetermined. San Juan Botanical Garden, San Juan, north-eastern Puerto Rico.

 

Bug, Spartocera sp. Grand Anse, south-western Grenada, Lesser Antilles.

 

Leaf-footed bug, (family Coreidae), probably Leptoglossus sp. Mount Hartman Bay, south-western Grenada, Lesser Antilles.

 

Assassin bug, probably Rocconota sp. Guanica State Forest, south-western Puerto Rico.

 

Phymatid ("ambush") bug, species undetermined. Florida, north-central Puerto Rico.

The shape of the front legs of this species to indicate that it is a predator of other insects.

 

Alydid bugs, Megalotomus sp., male and female. Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Bug, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Bug, species undetermined. San Juan Botanical Gardens, San Juan, north-eastern Puerto Rico.

This individual is playing dead by resembling a dead piece of bark.


Bug, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Membracid bugs, Umbonia crassicornis, females. San Juan Botanical Gardens, San Juan, north-eastern Puerto Rico.

 

    Cicadas, leafhoppers, and related species also have piercing mouthparts, and feed on the sap of diverse plants. The pre-adult stages of cicadas are subterranean, and feed on plants' roots. Adult males produce strident calls with their elytra, in order to attract mates.

 

Cicadas, species undetermined, female (top) and male. Central Mountain Range of the Dominican Republic, Hispaniola.

 


Cicada, Borencona aguadilla, male. Carite State Forest, east-central Puerto Rico.

Unlike some other cicadas, the call of this species is heard almost exclusively at night.

 

Cicada emerging from its last pupal stage, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Cicada nymph, probably Proarna hilaris. Guilarte State Forest, west-central Puerto Rico.

Cicada nymphs spend their lives underground, feeding on roots.

 

Cicada, species undetermined. Eggleston, south-central Dominica, Lesser Antilles.

 

Cicada, Chinaria sp., male. Puerto Escondido, south-western Dominican Republic, Hispaniola.

 

Cicada, Proarna hilaris, male. Juncos, east-central Puerto Rico.

 

Planthoppers (Family Fulgoroidea) Ormenaria rufifascia. Fort Buchanan, Guaynabo, north-eastern Puerto Rico.

 

Acalonid planthopper, species undetermined. Toro Negro State Forest, central Puerto Rico.

 

Leafhopper, Tylozygus sp. Guilarte State Forest, west-central Puerto Rico.

 

Order Hymenoptera: Wasps, Ants, and Bees

 

    These insects are mostly social, though many species are solitary. Examples of these are the most primitive among them, the suborder Symphita: the "sawflies". The larvae of these insects are free-living, are herbivores, and resemble caterpillars - indeed very similar to those of their distant relatives, the butterflies and moths. Females do not possess stingers, and adults can be both herbivores or predators of other insects.

 

Puerto Rican Krug's sawfly, Sericocera krugii. Eggs, larvae, and adult male. San Juan, north-eastern Puerto Rico.

Notice the similarity of the larvae with those of a lepidopteran caterpillar.

 

    More advanced hymenopterans possess a unique defensive and offensive apparatus, their stinger, which is actually a modified ovipositor, or egg-laying organ, attached to a venom gland located in the abdomen. Depending on the species, the sting's potency may be hardly noticeable, or might produce an excruciating and incapacitating pain.

 

    Having "invented" paper eons ago, many wasps make their free-hanging nests on the underside of boughs, leaves, and rocky ledges. In them they raise their larvae with a regurgitated pulp made out of insect and spider prey. Two genera are common, Polistes and the very similar Mischocyttarus, this last different from the former in having a long-petioled abdomen. Anyone who spends some time walking along a trail in most West Indian forests will eventually come face to face with a paper-wasp nest. That same person will know that the insects often take serious exception to an intrusion in their space. Being stung simultaneously by several individuals of the larger species is a perfect way to ruin a pleasant stroll.



Neotropical paper wasps, probably Mischocyttarus mexicanus.

First photograph: Sage Mountain National Park, west-central Tortola, British Virgin Islands.

Second photograph: Florida, central Puerto Rico.

 

Paper wasp, possibly Polistes americana, preying on a caterpillar. Guilarte State Forest, central Puerto Rico.

 

Paper wasps, probably Polistes sp. Vermont Nature Reserve, south-western Saint Vincent, Lesser Antilles.

 

Paper wasps, Polybia occidentalis. Their huge nest hangs from a thin branch in a mangrove swamp.

Mount Hartman National Park, south-western Grenada, Lesser Antilles.

 

A parasitic wasp, species undetermined, has laid its eggs on this caterpillar. After birth, the larvae will eventually eat the caterpillar alive from the inside out.

Camp Santiago, Salinas, south-eastern Puerto Rico.

 

    Some wasps and bees are solitary. The females build their nests underground or in dead wood, where they raise their larvae by feeding them insect or spider  prey (in the case of wasps) or honey (in the case of bees). Among Caribbean solitary wasps, tarantula hawks of the genus Pepsis are notorious for their large size and strikingly metallic colors. They are ferocious hunters of tarantulas and other large spiders, and show great ability in keeping just out of reach of their preys' deadly fangs. After harassing their victims until they are tired they sting them on their undersides. The paralyzed - but still living - spiders are placed in an underground chamber and a single egg is deposited on them. The larva hatches and then feeds to adulthood on the spider, which is being eaten from the inside out until only a hollow husk is left.

 

Tarantula hawk (family Pompilidae), Pepsis ruficornis, female. Near Rabo de Gato, south-western Dominican Republic, Hispaniola.

These solitary wasps hunt for spiders and other arthropods with which to feed their developing

larvae, hidden in subterranean nests. Many species of this genus possess a bright, metallic iridescence.



Tarantula hawk, Pepsis sp., female. Soufriere Botanical Garden, south-eastern Saint Lucia, Lesser Antilles.


Tarantula hawk, Pepsis ruficornis, female attacking a wolf spider. Guilarte State Forest, west-central Puerto Rico.

 

Pompilid wasp, probably Pepsis sp., sleeping on a leaf at night. Grand Etang National Park, central Grenada, Lesser Antilles.

 

Pompilid wasp, Prionyx thomae. Isabela, north-western Puerto Rico.

(Photograph courtesy of Dr. Javier Mercado).

 

Velvet ant, (family Mutillidae), probably Dasymutilla bouvieri, female. Rabo de Gato, south-western Dominican Republic, Hispaniola.

These "ants" are actually a kind of terrestrial, wingless wasps. Many species can deliver very painful stings.

 

Ichneumonid wasp, Ophion sp. Camp Santiago, Salinas, south-eastern Puerto Rico.

These wasps are parasites of other insects, laying their eggs on living hosts. The larvae hatch to feed on their victim.

 

Ichneumonid wasp, species undetermined. Del Este National Park, eastern Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Piero Fariselli).

 

Ichneumonid wasp, species undetermined. Guanica State Forest, south-western Puerto Rico.

(Photograph courtesy of Mr. Jose Rodriguez Molina).

 

Sand wasp, (family Crabronidae), Stictia signata. Susua State Forest, south-western Puerto Rico.

This species makes underground lairs in sandy areas, where its breeds its larvae.

 

Scoliid wasp, Campsomeris trifasciata, male. Sandy Point National Wildlife Refuge, south-western Saint Croix, United States Virgin Islands.

 

    Solitary bees in the region include the bumblebees of the genus Xylocopa. These last are frequently called "carpenter bees" for their habit of hollowing out nests in dead snags, where they build their nests. The mother accumulates a portion of pollen and honey in the grub's chamber, and then seals it off with a mixture of chewed wood and salive. Once it has metamorphosized, the new adult breaks open its imprisonment and leaves. The sexes of the common Antillean species are strikingly different in appearance. Females are jet black, while the rare and seldom seen males are rusty orange in color.

 

A female Xylocopa mordax peeks out of its hole excavated in a dead branch.

The males and females of these enormous bees are quite different, males being golden yellow.

Guayama, south-eastern Puerto Rico.

Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Apid bees, Anthophora tricolor. Florida, central Puerto Rico.

 

A small halictid bee, Habralitellus rufopanticis, collects nectar from a melastome flower, Leandra krugii.

Maricao State Forest, western Puerto Rico.

 

    The several families of ants comprise an early evolutionary offshoot of the wasp lineage. Some are herbivores, while others are omnivores or predators of other organisms. Like wasps, ants usually carry a sting, and most species live in colonies where individuals belong to one or another class of queen, drones, and sterile female workers and soldiers.

 

    Solenopsis fire ants are found in open, relatively dry microhabitats in mesic areas of the Antilles. Their name derives from the burning sensation caused by their stings and the reddish hues of most species. The sting of a single fire ant is by no means a major affair except for a hypersensitive or allergic person. The individual animals are very small, to begin with, and the amount of their venom is minute, as well. However, disturbing a fire ants' nest may mean courting disaster, since as many as hundreds of thousands of angry insects swarm over everything in their path, seeking the cause of their irritation. If the intruder is a small animal, it may easily be stung to death.

 

    Other West Indian species of ants, like the much larger, long-bodied and spindly-legged ponerines of the genus Odontomachus, establish their rather small colonies under rocks and downed, rotting trees. They are much less aggressive than fire ants but will attack when hard-pressed, biting down with their long, pincer-like jaws and then stinging painfully.

 

The nests of fire ants (Solenopsis sp). can be recognized by the mounds of soil that dot many grassy regions in the Neotropics.

The ants themselves are tiny, but possess a nasty sting.

They can represent a nuisance in agricultural areas, and may even constitute a moderate threat to human health.

Photographed in Aguas Buenas, central Puerto Rico.

 

Winged adult and worker of Solenopsis geminata fire ants.

Yauco, south-western Puerto Rico.

 

Ants, Odontomachus haematoda. These ponerines can deliver quite a painful sting.

Barrenspot, central Saint Croix, United States Virgin Islands.

 

Ants, species undetermined. Near Negril, western Jamaica.

 

Arboreal ants' nest. Barbecue Bottom, north-central Jamaica.

 

This nest of a Puerto Rican emerald hummingbird, Chlorostilbon maugeus, has been assailed by arboreal ants.

Unable to defend its chicks, the parent bird has abandoned them.

El Rosario, south-western Puerto Rico.

 

Carpenter ants, Camponotus kaura. Grand Colombier, Saint Bartholomew, Lesser Antilles.

(Photograph courtesy of Mr. Karl Questel).

 

Order Isoptera: Termites

 

    Termites are distant relatives of cockroaches which have adapted to digest cellulose (the main component of wood) with the help of protozoan symbiotes living in their guts. In terms of individuals, they are among the most abundant of terrestrial arthropods. Trillions of them live in Antillean forests, recycling dead wood and making it available again for use by other living organisms.

 


Nest of termites, Nasutitermes costalis. Mary Point, Virgin Islands National Park, Saint John, United States Virgin Islands.

The one on the left has been abandoned and is in a state of decay.
Termites are among the best known "social" insects on Earth, after hymetopterans (wasps, bees, and ants).
Distant relatives of cockroaches, their capability to digest cellulose, (with the help of protozoan symbionts that live in their guts)
places these insects among some of the most influential organisms in tropical forests' food chains,

by allowing them to digest enormous quantities of otherwise unusable organic matter. Only fungi,
as a whole, are more important on regards to the use of dead tissue in Earth's terrestrial ecosystems.

 

Nasutitermes costalis. Lajas, south-western Puerto Rico.

Soldiers of Nasutitermes have pointy heads. Indeed, the generic name translates to "termite with a nose".

From the tip of their "noses" they squirt a sticky, irritating fluid used in the defense of their colonies.

 

The galleries of Nasutitermes termites, made of wood pulp mixed with saliva, extend far away from their colonies.

These passages are used to travel under cover to and from sources of dead wood, like this dead tree stump.

Mary Point, Virgin Islands National Park, Saint John, United States Virgin Islands.

 

    Termites are social insects, and their groups, in a manner similar to wasps, and and bees, center around a queen, which is the only fertile female in the colony. The colonies of some species are the largest of any social insects, and sometimes contain millions of individuals.

 

Termites, species undetermined.

Near Rabo de Gato, south-western Dominican Republic, Hispaniola.

 

Order Lepidoptera: Butterflies and Moths

 

    This is the second largest order of insects, after Coleoptera (beetles). Each wing is covered by thousands to millions of microscopic scales that fit together among themselves like shingles on a roof. Each scale is of a single color but, to the naked human eye, the juxtaposition of the scales of different colors forms the overall pattern of the animals wings in the same way in which a mosaic is seen, as a whole. It is this trait which gives them its name, from the greek "lepido-pteron": "scaly wings".

 

    Their taxonomy is quite complex and about 126 families are known to date. Butterflies are grouped into two large superfamilies. Herperiodea contains the skippers and their allies, while all other (and the majority) of butterflies are placed in the Papilionoidea. Moths of the other hand, are far more abundant in terms of species, and are placed in about 42 superfamilies, themselves divided into many more families.

 

    Among insects, butterflies and their nocturnal relatives and counterparts, namely moths, attract attention for their frequently vivid colors. They are universally recognized and, except for species that resort to mimicry, they cannot be confused with anything else.

 

    Lepidopterans possess mouthparts modified for ingesting liquids by suction. A long proboscis is extended and inserted into flowers or wounds on plants in order to feed on nectar, tree sap, or juices of rotting fruits. The color patterns of their wings have a very peculiar origin.

 

    These scales are what is left on a hand as a powdery residue when one handles a butterfly or moth. As the animal grows older, and with the exertions of flying, the scales constantly fall off day by day. That is the reason why butterflies so often look "washed out" and discolored already a few weeks after emerging from the chrysalis. Indeed, the adult life of most butterflies and moths lasts only a few months, when not mere weeks, before their wings begin to literally fall apart.

 

    Although butterflies and moths are very similar, most butterflies can usually be told for what they are if one looks closely at their antennae. Those of butterflies are thin and end in a small club. The antennae of male moths are feather-like, while those of females are not, yet they never end in a club.

 

    Butterflies usually have a thin and pronounced "waist" that separates their thoraxes from their abdomens. Moths, on the other hand, lack this trait, their thoraxes and abdomens being nearly fused together without a clear demarcation between the two body parts. One clear exceptions to this rule are the skippers, (small butterflies of the family Hesperiidae), which indeed look like small moths in their general appearance.

 

    A less obvious difference between the two groups can be noticed when one looks at a resting individual of either kind. Butterflies will usually hold their wings closed, pressed together above the body, while moths will either hold them open and pressed against the substrate or will fold them backwards along and over their bodies. Again, exceptions exist, like that of the nymphalid butterflies of the genus Hamadryas, which will hold their wings open and flat against the tree trunks on which they usually alight.

 

    And finally, the least reliable difference: butterflies are creatures of the sunlight, while moths enter into their own at night. While no known West Indian butterfly is nocturnal, several moths in the region do indeed prefer the daylight for their feeding and mating. However, and most interestingly, most of the latter are species that mimic diurnal animals, sometimes taxonomically very different from them. The color and flight patterns of some Composia moths are somewhat similar to those of the poisonous Heliconius butterflies the inhabit many of the same forests. Other diurnal moths in the region look strikingly and, at first sight, alarmingly similar the fierce Polistes and Pepsis wasps with which the live in sympatry. And the small and green Aellopos sphynxid moths bear an astonishing resemblance to the tiny Caribbean Mellisuga and Chlorostilbon hummingbirds, as they nervously flit from one flower to another in search of nectar during the day (although, actually, the model for their mimicry seems to be a South American hummingbird, not found in the Antilles).

 

    There is no lack of mimics among nocturnal Caribbean moths, but these usually resemble inanimate objects, like the lichen or pieces of tree bark on which they usually rest during the day. However, some of the most attractive butterflies in these islands can spectacularly disappear from sight in a fraction of a second not only by coloration mimicry, but also through an interesting behavioral adaptation. This is especially noticeable in the case of some nymphalid butterflies like the, leafwings, silverwings, and orions of the genera Anaea, Siderone, Archaeoprepona,and Historis. These nervous and fast-flying insects will perch on a tree trunk or branch and will instantly close their wings. The upper surface of these are vividly colored in red, orange or blue, but their undersides are patterned like dead, dry leaves. The effect is that they seem to vanish into thin air, and then are almost impossible to spot unless they move, an almost certain way to remain undetected by at least some predators. The members of the also nymphalid genus Hamadryas exhibit a similar behavioral strategy though, in their case, their wings are somberly blotched in grays and browns, and are held flat against the tree trunks, which bark they sometimes mimic with almost total perfection.

 

    At the opposite end of this behavioral spectrum, some butterflies apparently want to be seen by their predators. These include the slow-flying and brilliantly colored danaiine and heliconiine nymphalid butterflies, like the Danaus monarchs and queens, the Lycorea tigers, the Eueides and Heliconius longwings, and the Agraulis fritillaries. These beautiful butterflies use their striking upper- and underwing patterns and their phlegmatic flights to advertise their distastefulness to potentially interested predators. Their repulsive flavors and dangerous toxicities are due to alkaloids that they incorporate directly from their larval food plants, usually Asclepias milkweeds and Passiflora vines. In turn, some of these butterflies are mimicked by others (like the pierids of the genus Dismorphia) which may or may not be toxic themselves, but which nonetheless derive some protection from the visual ruse.

 

The proboscis of a dusky swallowtail, Heraclides aristodemus aristodemus swallowtail butterfly can be seen under its head.

Guanica State Forest, south-western Puerto Rico.

 

A silverking butterfly, (Puerto Rican race, Archaeprepona demophoon ramorosum) feeds on trees sap with its long proboscis.

Camuy Caverns Park, Camuy, north-western Puerto Rico.

 

Black witch moth, Ascalapha odorata, male, feeding on the juices of fallen fruit.

San Juan north-eastern Puerto Rico.

 

    Although the butterfly and moth faunas of the Antilles are less varied than those of the nearby continents, they still are conspicuous members of the local insect fauna. A number of species are endemic to the Caribbean islands while many others are found in the American continents, as well.

 

Caterpillar, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Caterpillar mimicking tree bark, species undetermined. Cambalache State Forest, northern Puerto Rico.

 

Caterpillar, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

Caterpillar (species undetermined) feeding on the fruits of Gonzalagunia hirsuta.

Near Morne Anglais, southern Dominica, Lesser Antilles.

 

Caterpillar, species undetermined. Florida, central Puerto Rico.

 

Caterpillar, species undetermined. El Yunque National Forest, north-eastern Puerto Rico.

 

    Some of the largest and most beautiful butterflies in the world belong to the family Papilionidae. (In fact, Papilio is the genus in which Linnaeus placed all butterflies). Many species are commonly known as "swallowtails" for the extensions of their hind wings. The largest butterfly in the Americas is the giant swallowtail of Jamaica, Pterourus homerus. This huge and impressive insect is endangered due to habitat loss.

 

Androgeus swallowtail, Heraclides androgeus epidaurus, male, drinking from a puddle.

El Rosario, south-western Puerto Rico.

Many members of this family are commonly called "swallowtails",

for the elongated posterior tips of their back wings.

 

The huge caterpillar of Heraclides androgeus epidaurus.

Ciales, central Puerto Rico.

 

Machaonides swallowtail, Heraclides machaonides.

Second Picture taken at the Bahoruco National Park, south-western Dominican Republic, Hispaniola.

 

Dusky swallowtail, (nominate race, Heraclides aristodemus aristodemus).

Guanica State Forest, south-western Puerto Rico.

 

Pelaus swallowtail, (Hispaniolan and Puerto Rican race, Heraclides pelaus imerius), female.

Florida, central Puerto Rico.

 

Polydamas swallowtail (Puerto Rican race, Battus polydamas thyamus), caterpillar and adult.

First photograph: Crown Mountain, Saint Thomas, United States Virgin Islands.

Second photograph: Carite State Forest, east-central Puerto Rico.

 

    Satyrids are represented in the West Indies by a single genus endemic to the region: Calisto. These small butterflies, gray or reddish brown in color, can be seen fluttering close to the ground in all the Greater Antilles and some of the Bahamas. By far, most species are endemic to Hispaniola, which probably has several dozen species. A few are found in Cuba and the Bahamas, two inhabit the Puerto Rican bank, and there is a single species in Jamaica.

 


Puerto Rican calisto, Calisto nubila, females.

First photograph: Susua State Forest, south-western Puerto Rico.

Second photograph: Cambalache State Forest, northern Puerto Rico.

The entire genus of several dozen species is restricted to the Greater Antilles and the Bahamas, with most species

being endemic to Hispaniola. The family (Satiridae) is widespread in the Americas, especially in the Neotropics.

 

Confused calisto, Calisto confusa. Near Pie Pol, Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

 

Jamaican calisto, Calisto zangis. Blue and John Crow Mountains National Park, east-central Jamaica.

 

Yellow-barred calisto, Calisto archebates, male. Zapoten, south-western Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Pedro Genaro Rodriguez).

 

Calisto galli galli. Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Miguel Angel Landestoy).

 

    One of the families of large and colorful butterflies that is best represented in the Antilles is Nymphalidae. Many species are colorful, and members of the related genera Anaea, Archeoprepona, Historis, and Hypna are notoriously fast insects with powerful and bat-like flights. Some, like Colobura, Hamadryas, Marpesia, and the aforementioned Anaea and Archaeoprepona  are highly cryptic when they perch motionless, often folding their wings and showing their undersides that resemble dead leaves. The striking colors of the dorsal sides of their wings are only revealed when they fly.

 

    Some species like the monarch, Danaus plexippus (of the subfamily Danaiinae) are highly toxic to most predators. Their caterpillars feed on plants with high concentrations of alkaloids that the insects incorporate into their own tissues, thus making themselves unpalatable. They advertise their distastefulness with brilliant orange and black colors, an aposematic combination. Although continental populations of the monarch are famous for their migrations that span thousands of miles, West Indian populations are sedentary, and have indeed evolved into several local subspecies.

 

A male Puerto Rican crested anole (Anolis cristatellus cristatellus) attempts to devour the caterpillar of a Puerto Rican monarch butterfly (Danaus plexippus portoricensis), only to vomit it a few minutes later.

San Juan, north-eastern Puerto Rico.

 

    Many West Indian nymphalids do not feed on nectar, but rather ingest tree sap or the juices of rotting fruits. They sometimes aggregate whenever fallen fruits accumulate, or where woodpeckers or other animals have broken the bark of living trees.

 

Caterpillar and adult male of monarch butterflies (Puerto Rican race, Danaus plexippus portoricensis).

First two photographs: San Juan, north-eastern Puerto Rico.

Second photograph: Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Queen butterfly, (Lesser Antillean race, Danaus, gilippus xanthippus).

Mount Hartman National Park, south-western Grenada, Lesser Antilles.

 

Lesser false fritillary, (nominate race, Anetia briarea briarea), male.

Bahoruco National Park, south-western Dominican Republic, Hispaniola.

 

Jaeger's false fritillary, Anetia jaegeri, male.

Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Pedro Genaro Rodriguez)

 

Large tiger, Lycorea cleobaea cleobaea.

Near Rabo de Gato, south-western Dominican Republic, Hispaniola.

Members of Lycorea mimic the also distasteful heliconiid butterflies, Heliconius.

The kind of mimicry, called "Mullerian", involves two or more species that are both similarly

poisonous or toxic and similar in their appearance, and that usually live in sympatry.

In that way, they reinforce each another's effectiveness in dissuading predators.

 

Haitian sister, Adelpha gelania, male. Camuy Caverns Park, Camuy, north-western Puerto Rico.

 


Malaquite butterfly, Siproeta stelenes stelenes. Near Rabo de Gato, south-western Dominican Republic, Hispaniola.
Family Nymphalidae.

 

Siproeta stelenes stelenes. First photograph: Florida, central Puerto Rico.

Second photograph, Maricao State Forest, western Puerto Rico.

 

Saint Lucian mestra, Mestra cana, male. Mount Hartman National Park, south-western Grenada, Lesser Antilles.

 

Nymphalid butterfly, Asterocampa idyja. Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Miguel Angel Landestoy).

 

Cuban crescent, Anthanassa frisia frisia.

Near Polo, Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

 

Nymphalid butterfly, Doxocopa thoe, male.

Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Pedro Genaro Genaro Rodriguez).

 


Another Antillean nymphalid, the Puerto Rican leaf-wing, Anaea borinquenalis. Guanica State Forest, south-western Puerto Rico.
This cryptic species endemic to Puerto Rico resembles a dry leaf when it perches on a branch or on the ground.

However, when it opens its wings, it reveals their bright orange color.

 

Hispaniolan leaf-wing, Anaea troglodyta, female.

Rabo de Gato, south-western Dominican Republic, Hispaniola.

 

Jamaican leaf-wing, Anaea portia, female. Sheffield, western Jamaica.

 

False neptis, Archmestra teleboas.

Near Rabo de Gato, south-western Dominican Republic, Hispaniola.

The genus is monotypic, and endemic to the Hispaniolan insular bank.

 

White peacock (nominate race, Anartia jatrophae jatrophae).

Mount Hartman National Park, south-western Grenada, Lesser Antilles.

 

White peacock (Puerto Rican race, Anartia jatrophae semifusca).

Caño Tiburones Nature Reserve, northern Puerto Rico.

 

Tropical buckeye, Junonia genoveva, male. Camp Santiago, Salinas, south-eastern Puerto Rico.

 

Anartia lytraea. Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Mosaic butterfly, Colobura dirce wolcotti.

First photograph: Florida, central Puerto Rico.

Second photograph: El Yunque National Forest, north-eastern Puerto Rico.

 

Mexican fritillary (Puerto Rican race, Euptoieta hegesia watsoni).

Guanica State Forest, south-western Puerto Rico.

 

Puerto Rican harlequin, Atlantea tulita, eggs, caterpillar, and adult. Guajataca, Quebradillas, north-western Puerto Rico.

Each of the four members of this genus is endemic to an Greater Antillean bank.


Red rim, Biblis hyperia, takes off from a leaf, and then alights on another.

Guanica State Forest, south-western Puerto Rico.

Two Eunica tatila tatilisca (also nymphalids) accompany the second individual.

 

Florida purplewing, (Antillean race, Eunica tatila tatilisca), female. Ebano Verde Scientific Reserve, central Dominican Republic, Hispaniola.

(Photograph courtesy of Mr. Miguel Angel Landestoy).

 

Dingy purplewing, Eunica monima.

First two photographs: males. Guanica State Forest, south-western Puerto Rico.

Last photograph: female, Puerto Escondido, south-western Dominican Republic, Hispaniola.

 

Silverking, Archaeoprepona demophoon (Hispaniolan subspecies, A. d. amphitoe).

National Botanical Garden, Santo Domingo, southern Hispaniola.

A leaf-mimic when at rest, the upper surface of the wings have iridescent blue markings on a blackish brown background.

 

And this is the Puerto Rican race of the silver king, A. d. ramorosum.

Susua State Forest, south-western Puerto Rico.


The largest nymphalid in the West Indies: the orion, Historis odius odius.

Near Pie Pol, Bahoruco Mountains, south-western Domini