Butterflies

Papilionoidea (= Rhopalocera). Spanish: Mariposas. Portuguese: Borboletas. Nahuatl: Papalomeh, sing, papalotl (Mexico). Quechua: Pillpinto, ccori kente. Tupi-Guaraní: Panamá.

Truly, the resplendent colors and airy flight of butterflies give the Neotropical environment much of its charm and beauty, for these insects are nowhere else so abundant and diverse. The fauna is the largest for any zoogeographic region (an estimated 7,000—9,000 species or nearly 50%; Legg 1978, Robbins 1982) and contains some unique types that are among the most attractive in the world. The wings of morphos, for example, have been used to make jewelry and the famous "art deco" serving trays and montages sold as art pieces as an early tradition in the tourist centers of Brazil and other parts of South America. In fact, the demand for butterfly artwork is so great that an economically significant cottage industry has prospered for a long time (Barrett 1902) in some Latin American countries. It is especially well developed in Brazil and Peru, where organized marketing chains exist. In Peru, this begins with the jungle peasant collectors, who peddle the raw specimens to traders in Tingo María, Satipo, and Pucallpa, who in turn ship them to craftsmen and merchants in Lima where they are mounted and finally displayed in the tourist shops. Most are sold to visitors as curios; few become material for collectors or scientists.

The trade has some strange beginnings in the practices of earlier inhabitants. Collecting butterflies for sale in Europe and North America provided the only real source of survival for many French libérés from the infamous Devil's Island penal colony. These men (Norris 1955), some famous, such as René Belbenoit, expositeur of the abominable conditions in the prison (Dry Guillotine), and his famous fanciful counterpart Henri Charrière (Papillon), were condemned to spend their remaining days on the lie Royal and eked out an existence by selling morphos and other valued species to middlemen.

The possible deleterious effects of such resource exploitation have only been partly assessed (Pyle 1976). Some entomologists (Carvalho and Mielke 1972) feel that little harm is done, since, for the most part, individuals that are harvested are males that have already mated or females that have laid their eggs. It is believed that natural enemies, such as lizards (Ehrlich and Ehrlich 1982) and birds, are more effective in controlling populations. Others, however, are of the opinion that considerable genetic damage to natural butterfly populations may ensue from commercial collecting (Owen 1974).

The attitudes of ancient Mexicans were less pecuniary. They stylized many species in their religious art (Franco 1961, de la Maza 1976) and recognized a butterfly (Papilio multicaudata) as Xochiquetzal, Goddess of Flowers. Numerous images of butterflies are included in the frescoes of the temple of Tlalocân in Teotihuacân near Mexico City (Beutelspacher 1976). Butterflies were thought to be the reincarnated souls of dead warriors returned to earth. The ancient Aztec name for the butterfly, papalotl, persists in the Mexican language of today in place-names (Papalotepec, Mexico) and in other forms; Indian children still a kite amapapalotl (paper butterfly) (Hoffman 1918).

Modern scientific study of butterflies is current'y very active with regard to the Neotropical fauna; the early admonition of Yi. W. Bates (1892: 353) seems at last to have been realized: "the study of butterflies— creatures selected as the types of airlines and frivolity—instead of being despised, ^¡11 some day be valued as one of the most important branches of biological science." Indeed, active research on Neotropical butterflies is having an impact in many areas of ecology (Gilbert and Singer 1975, Young 1980a), evolution (Benson 1971), genetics and mimicry (Turner 1977), cytology (Wesley and Emmel 1975), physiology (Swihart 1972), and biogeography (Brown 1981) and to other aspects of general biology and science.

Nevertheless, many widespread phenomena are still poorly understood. The causes and patterns of migration of enormous numbers of many species, for example, remain unknown (Welling 1959). Clouds of butterflies, often yellow pierids, were described by early chroniclers in Amazonia and the Caribbean where they are best seen. Darwin (1962) tells of a time when the Beagle was some miles off the mouth of La Plata: "vast numbers of butterflies, in bands or flocks of countless myriads, extended as far as the eye could range. Even by the aid of a telescope it was not possible to see a space free from butterflies. The seamen cried out 'it was snowing butterflies,' and such in fact was the appearance." Such migrations are even recognized by a special word in the widespread Tupi language, panapana.

The purpose of "puddling," the habitual practice of many butterflies (mostly yellows and swallowtails and almost always the males) to siphon liquids from moist •oil, is also still a puzzle (Boggs and Jackson | 1991). Dense groups of individuals thus disposed along tropical lowland river-i> often at spots where humans laun der their clothes or bathe, are a common sight. The insects rest side by side, with the axis of their bodies aligned, and suck up with their tongues interstitial fluids from sand and mud, pumping it through their bodies until it at times fairly squirts from the anus. Some important nutritional element could be removed in the process, but there may be other reasons for the behavior (see Lepidoptera, above). Children delight in chasing through these aggregations, sending throngs of butterflies into kaleidoscopic aerial dances, from which individuals often break away into "follow the leader" games (Collenette 1928).

Many butterflies also are fond of feeding on bird droppings (Young 1984), sweet-smelling, highly odoriferous rotting fruits, and decaying fungal growths on soupy sap flows (Young 19806). Some unknown special nutritive requirement is probably satisfied by these habits (ibid.). A number of species have also been observed drinking tears from the eyes of croco-dilians and turtles for some unexplained reason (Lamas 1986).

Sound is perceived by many species and is probably involved with territoriality and protection. Wing sacs at the base of the wing veins on the underside react to sonic stimuli (Swihart 1967).

The larvae of many Neotropical butterflies, especially among the metalmarks (Ly-caenidae, Riodininae) (Callaghan 1977), and blues (Lycaeninae) have evolved symbiotic partnerships with ants (Hinton 1951, Pierce 1987). They secrete substances that are sought by the ants, which jealously protect the larvae by aggressively biting or stinging any creature attempting to molest them. They may even protect the larvae by building shelters for them. The true nature of these associations remains largely unknown, although a considerable amount has been learned about some species (Malicky 1970). Many butterfly caterpillars are spiny (Nymphalidae), but the hairs borne on the spines are not urticating. Such larvae often have a long pair of spines arising on the head to distinguish them from similar spiny moth larvae (Saturniidae) whose heads are always hornless.

Another perplexing question about Neotropical butterfly biology is how mimetic species find their own kind for mating among the many almost identically appearing species in an area. It has been discovered that the wing patterns of many butterflies appear very different under ultraviolet illumination than under normal light (Mazokhin-Porshnyakov 1957) and constitute sexual signals. The eyes of the butterflies themselves have been found to be especially sensitive to the ultraviolet portion of the light spectrum, which explains how many mimetic species may distinguish between wing patterns that appear the same to us or to birds, their chief predators (Remington 1973). Odor clues also play an important part in sexual recognition (Pliske 1975).

A still further puzzling habit of some butterflies is their attraction to swarms of army ants (Eciton). Some observations of this phenomenon suggest that the butterflies drink protein and sugar-rich fluids from the droppings of ant birds that also flock to such swarms (Lamas 1983, Ray and Andrews 1980).

The richest area for butterflies is the Neotropics. There are four times as many butterfly species in Panama than in the entire Malaysian Archipelago and about twice as many skippers. The Tambopata Forest Reserve near Puerto Maldonado in southeastern Peru hosts some 1,200 species, the largest number for any comparable site in the world (Lamas pers. comm.). Butterflies are the best-known large group of insects, and it is possible to identify most to species using the many works available (Lamas 1977, 1978). Some excellent popular guides to the adults have been published (e.g., d'Abrera 1981, 1984a, 19846, 1987a, 19876, 1989; Barcant 1970; De Vries 1987; Riley 1975).

References

Barcant, M. 1970. Butterflies of Trinidad and Tobago. Collins, London.

Barrett, O. W. 1902. Cheap tropical butter flies. Entomol. News 13: 239-240.

Bates, H. W. 1892 [1863], The naturalist on the River Amazons. Murray, London.

Benson, W. W. 1971. Evidence for the evolution of unpalatability through kin selection in the Heliconiinae (Lepidoptera). Amer. Nat. 105-213-226.

Beutelspacher, C. R. 1976. La diosa Xochi-quetzal. Soc. Mexicana Lepidop. Bol. lnf 2-1-3. ' '

Boggs, C. L., and L. A. Jackson. 1991. Mud puddling by butterflies is not a simple matter Ecol. Entomol. 16: 123-127.

Brown, Jr, K. S. 1981. Biogeography and evolution of Neotropical butterflies. In T. C. Whitmore and G. R. Prance, eds., Biogeography and Quaternary history in tropical America. Oxford Univ. Press, Oxford.

Callaghan, C. J. 1977. Studies on restinga butterflies. 1. Life cycle and immature biology of Menander felsina (Riodinidae), a myr-mecophilous metalmark. J. Lepidop. Soc. 31-173-182.

Carvalho, J. C. M., and O. H. H. Mielke. 1972. The trade of butterfly wings in Brazil and its effects upon the survival of the species. 9th Int. Cong. Entomol. (Moscow) ): 486-488.

Collenette, C. L. 1928. Gatherings of butterflies on damp sand, with notes on the attraction of moths to human perspiration. Entomol. Soc. London Trans. 76: 400-407.

d'Abrera, B. 1981. Butterflies of the Neotropical Region. Pt. 1. Papilionidae & Pieridae. Lansdowne, East Melbourne, Australia.

d'Abrera, B. 1984a. Butterflies of the Neotropical Region. Pt. 11. Danaidae, Ithomiidae, Heliconidae, Morphidae. Hill House, Victoria, Australia.

d'Abrera, B. 19846. Butterflies of South America. Hill House, Victoria, Australia.

d'Abrera, B. 1987a. Butterflies of the Neotropical Region. Pt. III. Brassolidae, Acraei-dae and Nymphalidae—partim. Hill House, Victoria, Australia.

d'Abrera, B. 19876. Butterflies of the Neotropical Region. Pt. IV. Nymphalidae— partim. Hill House, Victoria, Australia.

d'Abrera, B. 1989. Butterflies of the Neotropical Region. Pt. V. Nymphalidae—partim. Hill House, Victoria, Australia.

Darwin, C. 1962. The voyage of the Beagle.

jjoubleday and Amer. Mus. Nat. Hist., New York.

r iA Maza, R. 1976. La mariposa y sus estilizaciones en las culturas Teotihuacana (200 a 750 d.c.) y Azteca (1324 a 1521 d.c.).

hg Vries, R J- 1987. Butterflies of Costa Rica i, jlid their natural history: Papilionidae, Pieri-dae, Nymphalidae. Princeton Univ. Press, Princeton.

EHRI-ich> P AND EHrl1ch- 1982.

Lizard prédation on tropical butterflies. J. Lepidop. Soc. 36: 148-152.

ffeanco, J- L. 1961. Representaciones de la mariposa en Mesoamérica. El México Antiguo 9: 195-244.

Gilbert, L. E., and M. C. Singer. 1975. Butterfly ecology. Ann. Rev. Ecol. Syst. 6: 365-397.

hlnton, H. E. 1951. Myrmecophilous Lycaeni-dae and other Lepidoptera—A summary. Proc. Trans. So. London Entomol. Nat. Hist. Soc. (1949-50): 111-175.

Hoffman, C. C. 1918. Las mariposas entre los antiguos mexicanos. Cosmos 1.

Lamas, G. 1977. Bibliografía de catálogos y listas regionales de mariposas (Rhopalocera) de América Latina. Soc. Mexicana Lepidop. Publ. Esp. Pp. 1-44.

Lamas, G. 1978. Adiciones a la bibliografía de catálogos y listas regionales de mariposas de América Latina (Rhopalocera). Soc. Mexicana Lepidop. Bol. Inf. 4(5): 1-14.

Lamas, G. 1983. Mariposas atraídas por hormigas legionarias en la Reserva de Tambopata, Perú. Rev. Soc. Mexicana Lepidop. 8(2): 49-51.

Lamas, G. 1986. Drinking crocodile tears. Antenna 10(4): 162.

Legg, G. 1978. A note on the diversity of world Lepidoptera. Biol. J. Linnean Soc. 10: 543-347.

Maucky, H. 1970. New aspects on the association between lycaenid larvae (Lycaenidae) and ants (Formicidae, Hymenoptera). J. Lepidop. Soc. 24: 190-202.

Mazokhin-Porshnyakov, G. A. 1957. Reflecting properties of butterfly wings and role of ultra-violet rays in the vision of insects. Biophysics 2: 352-362.

Norms, W. E. 1955. Dawn adventure. Chambers Journal, London.

|Ow£N, D. F. 1974. Trade threat to butterflies. Oryx 12: 479-483.

■CE, N. E. 1987. The evolution and bio-geography of associations between lycaenid butterflies and ants. Oxford Surv. Evol. Biol.

Pliske, T. E. 1975. Courtship behavior and use of chemical communication by males of certain species of ithomiine butterflies (Nymphalidae: Lepidoptera). Entomol. Soc. Amer. Ann. 68: 935-942.

Pyle, R. M. 1976. The ecogeographic basis for lepidoptera conservation. Ph.D. diss., Yale Univ., New Haven.

Ray, T. S., and C. C. Andrews. 1980. Ant butterflies: Butterflies that follow army ants to feed on antbird droppings. Science 210: 1147-1148.

Remington, C. L. 1973. Ultraviolet reflectance in mimicry and sexual signals in the Lepidoptera. New York Entomol. Soc. J. 81: 124.

Riley, N. D. 1975. A field guide to the butterflies of the West Indies. Collins, London.

Robbins, R. K. 1982. How many butterfly species? Lepidop. Soc. News (1982): 40-41.

Swihart, S. L. 1967. Hearing in butterflies (Nymphalidae: Heliconius, Ageronia). J. Ins. Physiol. 13: 469-476.

Swihart, S. L. 1972. Modelling the butterfly visual pathway. J. Ins. Physiol. 18: 1915-1928.

Turner, J. R. G. 1977. Butterfly mimicry: The genetical evolution of an adaptation. Evol. Biol. 10: 1636-206.

Welling, E. C. 1959. Notes on butterfly migrations in the peninsula of Yucatán. J. Lepidop. Soc. 13: 62-64.

Wesley, D. J., and T. C. Emmel. 1975. The chromosomes of Neotropical butterflies from Trinidad and Tobago. Biotropica 7: 24-31.

Young, A. M. 1980a. Evolutionary responses by butterflies to patchy spatial distributions of resources in tropical environments. Acta Biotheor. 29: 37-64.

Young, A. M. 19806. The interaction of predators and "eyespot butterflies" feeding on rotting fruits and soupy fungi in tropical forests; variations on a theme developed by the Muyshondts and Arthur M. Shapiro. Entomol. Rec. J. Var. 92: 63-69.

Young, A. M. 1984. Ithomiine butterflies associated with non-antbird droppings in a Costa Rican tropical rain forest. J. Lepidop. Soc. 38: 61-63.

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