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Lepidoptera.-- Entomophagous behavior has developed in a large number of families of Lepidoptera, which has predominantly phytophagous species. Early summaries were presented by Brues (1936), who also discussed the manner by which the change from plant to animal feeding probably occurred. Balduf (1931) reviewed information relating to predatory and parasitic species and later (Balduf 1938) gave an interesting account of the probable origin and rise of the entomophagous habit among lepidopterous larvae. Entomophagous Lepidoptera are thought to be derived mainly from ancestors that fed on the plant cortex and lichens. Entomophagy in this order is manifested in several ways, by cannibalism, occasional predation, habitual predation and parasitism. Cannibalism is an occasional diversion from the normal plant-feeding habit which results mainly from hunger of thirst. Occasional predation occurs in two forms, (1) that found among caterpillars of plant-feeding behavior, which sometimes attack insects beyond their own species, and (2) among scavengers, which may prey on other species occupying the same habitat. Two forms of habitual predators are, (1) several families with larvae that feed on Homoptera, and (2) a few species of Lycaenidae which is phytophagous in early larval instars, but later changes to predation.
Among butterflies, predators are found only in the Lycaenidae, while among moths it occurs in a large number of families and varies in extent of its development from occasional or chance predation to obligatory predation. The highest development of entomophagous behavior is found in Epipyropidae, some species being obligate external parasitoids, and individuals of all species being limited to a single host during larval development. True parasitism by members of other families is rare to find (Clausen 1940/1962).
The great majority of the hosts of predaceous Lepidoptera are in the Homoptera: Aphididae, Cicadellidae, Fulgoridae and Coccidae. Balduf considered that limitation to these groups was a result of their greater availability as compared with other phytophagous insects. They occur generally in all sections and upon a wide variety of plants, and are frequently present in abundance, and are soft in body and small.
The large order of Lepidoptera includes moths and butterflies. It has some of the most species of any order of animals. In the insects it includes the moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies are found almost everywhere. The name is derived from Ancient Greek λεπίδος (scale) and aptera (wing). Including over 160,050 described species, in 127 families and 46 superfamilies, the Lepidoptera show many variations of the basic body structure, which have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest that the order may have more species and is among the four largest, most successful orders, along with the Hymenoptera, Diptera, and the Coleoptera.
Most species are distinguished by being covered in scales, having two large compound eyes, and an elongated mouthpart called a proboscis. Almost all species have membranous wings, except for a few that have cross vein wings. The larvae are referred to as caterpillars and are different in shape, having a cylindrical body with a well-developed head, mandible mouthparts, and from 011 pairs of legs.
Over millions of years, there has evolved a large range of wing patterns and colorations ranging from colorless moths to the brightly colored and complex-patterned butterflies. Because of this Lepidoptera is the most recognized and popular of insect orders. Many species of the order are of economic value because of their importance as pollinators for the silk they produce.
The word Lepidoptera derives from the Latin word for "scaly wing" and from the Ancient Greek λεπίς (lepis) meaning scale and πτερόν (pteron) meaning wing. Sometimes the term Rhopalocera is used to group the species that are butterflies.
The origins of the common names of many species vary. The word butterfly is from Old English buttorfleoge. The species of Heterocera are called moths. The origins of the word moth may be related to Old English maπa meaning "maggot."
Lepidoptera are very successful insects, and are found on all continents, except the Antarctic. They inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus. They are mainly associated with higher plants, especially angiosperms. The northern-most of butterflies and moths is the Arctic Apollo (Parnassius arcticus) that ranges in the Arctic Circle in northeastern Yakutia, at an altitude of 1505 meters above sea level. In the Himalayas, some Apollo species such as Parnassius epaphus, have been found to occur up to an altitude of 6,000 meters above sea level.
Out of the more than 180,050 described species it is thought that 9 percent are butterflies and skippers with moths making up the rest. The vast majority of Lepidoptera are to be found in the tropics but a substantial biodiversity occurs on each continent, with some 11,300 species found in North America, and over 10,000 species reported from Australia.
Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from only a few millimeters long, to those with a wingspan of many inches, such as the Monarch butterfly. There are many variations of the basic body structure, which have evolved over time.
Lepidopterans like all Holometabola, undergo complete metamorphosis, going through a four-stage life cycle: egg; larva / caterpillar; pupa / chrysalis; and imago (plural: imagines) / adult. The morphological characteristics that distinguish the order Lepidoptera from other insect orders are as follows:
There are large compound eyes and mouthparts, the latter usually being modified into a proboscis; scales cover the external surface of the body and appendages; the prothorax is usually reduced; there are 2 pairs of wings, which have very few cross veins in most species; the posterior abdominal segments are modified for reproduction, and cerci are absent; the larvae are caterpillars and eruciform in shape with a prominent head and mandibles; prolegs are either absent or 8-10 in number; the pupae are primarily adecticous and obtect, but sometimes decticous.
The larvae, caterpillars, have a sclerotized head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections. There are 3 pairs of true legs, and additional prolegs (up to 5 pairs). Most caterpillars are herbivores, but some are carnivores of other insects. Larvae are the feeding and growing stages that undergo hormone-induced ecdysis, becoming larger with each instar. . The pupa is referred to as a chrysalis. There are functional mandibles and appendages that are fused or glued to the body in most species. The larvae of many species either construct a cocoon with a casing of silk, or pupate inside them or they will pupate in a subterranean cell.
Adults have two pairs of membranous wings that are usually completely covered by minute scales. Sometimes the wings are reduced or absent in the female but not the male. Antennae are conspicuous. In moths, the males often have more feathery antennae than females. The mouthparts of adults include a prominent proboscis formed from maxillary structures, and are adapted for sucking nectar. Some species have reduced mouthparts (some species do not feed as adults), and others have them modified to pierce and suck blood or plant fluids. Mandibles in adults are absent in all except the Micropterigidae that have chewing mouthparts. Adults also have two immobile, compound eyes, and, only two simple eyes or ocelli, which may be reduced. The three segments of the thorax are fused and consist of fixed sclerites. The wings are attached to the thoracic segments and are functionally dipterous due to a number of wing-locking mechanisms. Some species have flightless females with reduced wings. Abdominal segments 710 or 810 form the external genitalia. These genitalia are complex and diagnostic for most families and also serve in family identification. The more advanced families have the abdomen connected to the thorax by muscles with projections from the second abdominal sternite. Paired hearing organs at the base of the abdomen occur in the Pyraloidea and Geometroidea. Males have glandular and expandable organs that look like hairbrushes or tufts, or as thin-walled, eversible sacs.
The wings, head parts of the thorax and abdomen are covered with tiny scales. Most scales are lamellar, or blade-like and attached with a pedicel, while in other groups they may be hair-like or specialized into secondary sexual characteristics. The lumen or surface of the lamella has a complex structure. It shows color either due to the pigment colors contained within or due to its three-dimensional structure. Scales provide a number of functions, including insulation, thermoregulation, or aiding gliding flight. The most important function is the large diversity of vivid or indistinct patterns they provide, which gives camouflage, mimicry, and attracts mates.
Species that undergo complete metamorphosis have a life cycle that usually includes an egg, larva, pupa, and an imago or adult. The larvae are referred to as caterpillars, and the pupa of moths is called a cocoons and that of butterflies a chrysalis.
Mating begins with mate attraction, regularly using visual stimuli, especially in diurnal species such as most butterflies. However, most nocturnal female species, including most moths, use pheromones to attract males, sometimes from great distances. Some have acoustic courtship, or attract mates using sound or vibration.
Sexual reproduction is the most common as is egg laying. But some species give to live birth through ovoviviparity. There are differences in egg laying and the number of eggs laid. Some species with polyphagous larvae drop their eggs in flight, while most will lay their eggs near or on the host plant that the larvae feed on. The number of eggs laid varies from only a few to several thousand.
The larvae appear very different from the adults and come in a variety of shapes and sizes. They are characterized by an elongated body with 011 pairs of abdominal legs (usually 8) and apical crochets. Most have a well-developed head with mandibles. The larvae eat every part of the plant, and are considered pests. Some species lay their eggs on the fruit and others on clothing or fur (e.g., Tineola bisselliella, the common clothes moth). In Hawaii there is a carnivorous larva that feeds on flies. Some species are carnivorous and others are even parasitic. The larvae develop quickly with several generations per year. Nevertheless, some species may take up to 3 years to develop.
There are about 5-7 instars, or molts, regulated by certain hormones like prothoracicotropic hormone that stimulates the production of ecdysone, that initiates molting. Then, the larva puparium, which is the hardened cuticle of the last larval instar, develops into the pupa. The pupa may be covered with silk and attached with many different types of debris or nothing at all. The time it takes for pupae to emerge varies among species. The adult emerges from the pupa either by using abdominal hooks or a projection from the head.
Most Lepidoptera are terrestrial, but many species of Pyralidae are aquatic with all stages except the adult living in water. Many species from other families such as Arctiidae, Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae, Cossidae and Sphingidae are aquatic or semi-aquatic.
Locomotion in most species is flight. However, some species merely glide. Flight is either hovering, or a forward or backward motion.
Navigation is used by migrating species. Butterflies, that have more species that migrate, navigate using time compensated sun compasses. They see polarized light and therefore orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is thought to be especially important. Most migratory butterflies are those that occur in semi-arid areas where breeding seasons are short. The life histories of their host plants also influence them. Landscapes may also be used for navigation, such as coastal lines, mountains, roads etc. When flying over oceans flight direction is more accurate if the landscape on the coast is still visible. Moths also show navigation by using the Earth's magnetic field. Some months can correct a course with changing winds, and prefer flying with favorable winds. Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field.
Moths tend to circle artificial lights repeatedly, which suggests that these species use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, they can fly in a straight line. Celestial objects are so far away, that even after traveling great distances, the change in angle between the moth and the light source is negligible. Also, the moon is always in the upper part of the visual field. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth attempts to correct by turning toward the light, causing airborne moths to fly downwards. Moths also may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety and are thus inclined to circle ambient objects in the Mach band region.
Migration is usually seasonal, moving to escape dry seasons or other adverse conditions. Most species that migrate are butterflies, varying from short to over long distances. Moths also undergo migrations. . Some Neotropical species have population explosions and massive migrations. In Central America, the first population migrations may begin in July and early August and, depending on the year, may be very massive, continuing for as long as five months.
Lepidoptera are soft bodied, fragile and quite defenseless, and the immature stages move slowly or are immobile. Therefore, all stages are exposed to predation. Birds, lizards, amphibians, dragonflies and spiders, besides others, predate upon adult Lepidoptera. Caterpillars and pupae may be prey, not only to birds but invertebrate predators, small mammals, as well as fungi and bacteria. Parasitoid and parasitic wasps and flies may lay eggs in the caterpillar that would eventually kill it as they hatch inside its body and eat its tissues. Insectivorous birds are probably the principal predators as they serve an important food to many insectivorous birds.
An evolutionary interaction occurs between predator and prey species. Lepidoptera have developed a number of strategies for defense and protections, which include evolution of morphological characters, changes in ecological life-style and in behavior. These include aposematism, mimicry, and camouflage, development of threat patterns and displays.
Only a few birds, such as the nightjars, hunt nocturnal Lepidoptera and their main enemy are bats. Numerous evolutionary adaptations of moths occur to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidoptera eggs also suffer predation. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic of larvae of the same species.
Some species are poisonous to predators, such as the Monarch butterfly in America, as well as Papilio antimachus and the birdwings, the largest butterflies in Africa and Asia respectively. The toxicity is obtained plant chemicals. However, some species manufacture their own toxins. A predator that has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future.
Toxic butterflies and larvae usually have bright colors, striking patterns as an indicator to predators about their toxicity. This phenomenon is called aposematism. Other caterpillars emit bad odors to ward off predators. Some caterpillars, especially members of Papilionidae, have an osmeterium, a Y-shaped gland that projects and which is found in the prothoracic region of the larvae. When threatened, the caterpillar emits unpleasant odors from the organ to ward off the predators.
Camouflage and mimicry are also important for defense. Some species blend into their surroundings, making them difficult to be seen by predators. Caterpillars can be shades of green that matches the host plant. Others look like inedible objects, such as twigs or leaves. The larvae of some species, such as the Common Mormon (Papilio polytes) and the Western Tiger Swallowtail look like bird droppings. For example, adult Sesiidae species (also called clearwing moths) have a general appearance that resembles a wasp or hornet cause a Batesian mimicry.
Eyespots are a type of automimicry used by some lepidopterans. In butterflies, the spots are made up of concentric rings of scales of different colors.
Little is known of ancestral Lepidoptera species because so few fossils have been found. The earliest known fossil, Archaeolepis mane is from the Jurassic period, about . The fossil consists of a pair of wings with scales that are similar to the wing venation pattern found in Trichoptera (caddisflies). 2 other sets of Jurassic Lepidopteran fossils have been found, and 13 sets from the Cretaceous period. The best-preserved fossil is the Eocene Prodryas persephone from the Florissant Fossil Beds.
Lepidoptera are not common in areas where fossilization can occur, such as lakes and ponds, and their juvenile stage has only the hard head capsule that might be preserved. Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines are also seen in fossil leaves.. The earliest fossil is Archaeolepis mane from the Jurassic, about in Britain. It consists of wings and shows scales with parallel grooves and the characteristic wing venation pattern shared with Trichoptera. Only two more sets of Jurassic fossils have been found, and 13 sets in the Cretaceous. From there, many more fossils are found from the Tertiary, and especially Eocene Baltic amber
Most Lepidoptera are able to pollinate flowers. The adults feed on the nectar inside flowers, using their proboscis to reach the nectar. In the process, the adult brushes against the flower's stamen, on which the flower's reproductive pollen is stored. The pollen is then transferred to the adult that flies to the next flower to feed, where the pollen germinates and fertilizes the ova.
The larvae of Bombyx mori are the silkworms. They construct cocoons out of silk, which has been important economically. The species Bombyx mori has been domesticated to where it is completely dependent on humans for survival. However, the species Bombyx mandarina, or "Wild Silkmoth," lives and produces silk naturally.
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