川 TTERFLIES PhilipJ. DeVries Center for BiodiversityStudies, Milwaukee PublicMuseum Overview of Butterfly Taxonomic Diversity BUTTERFLIES LIKELY REPRESENT the most familiar I1. Early Stages and Host Relationships and best known group of all insects. Within the II1. Butterfly-Ant Symbioses context of human society, butterflies serve as center ﾂ IV. Butterfly Mimicry and Diversity pieces in educational media, they are used extensively V. GeographicalPatterns of Butterfly Diversity in the arts, including nature and commercial advertis ﾂ V1. Butterfly Diversity and Habitat Destruction ing, and they are used as symbols for religious and social groups. Within the field of biology, studies on butterflies have been fundamental to the development GLOSSARY of biogeography, behavior, coevolution, conservation, development, ecological genetics, evolution,global warming, mimicry, ﾂ aposematic Describing an organism that is rendered population ecology, sexual selec tion, speciation, symbiotic associations, and systemat- less susceptible to predation by advertising its obvi ﾂ ics. In summary, butterflies have been important to ous unpalatability. Batesian mimicry A form of mimicry in which the how we perceive biodiversity. target organism is rendered less susceptible to preda ﾂ tion by its resemblancein morphologyor coloration to a different species that is unpalatable. 1.OVERVIEW OF BUTTERFLY cryptic Describing an organism that is concealed or TAXONOMICDIVERSITY obscured by the similarity of its appearance to the surrounding environment. Although butterflies may be the best known group of Mullenan mimicry A form of mimicry in which two insects, our understandingof their taxonomic diversity or more unpalatable species resemble each other, has two fundamentalweaknesses. The first regards the with the effect that predators are more likely to avoid recent declineof professional butterflytaxonomists and any species with this appearance. specieslevel revisions in the past 50 years. This requires myrmecophilyAbility to form symbiotic associations most species diversityestimates (from family to genus) with ants. to be derived from a literature that is out-0f-date. The vibratory papillae Mobile, grooved, rod-1ikeappend ﾂ second is that the number of families and subfamilies of ages arising from the distal edge of the first thoracic butterfliesvaries among classificationsbecause, despite segment, used for communicating. current interest in phylogenetic systematic methodsand analyses,the relationships within the major groups are EC ncydope mO lイ@vc ヴY 山でmel 卜 。 opyngh byACl而ade ヒミpr orights uで lionin Or血 an reserved 559 560 . BUTTERFLIES unresolved, particularlywithin Nymphalidaeand Ly- nectar, and males drink at wet soil (Fig. 1); caterpillars caenidae. Such is the state of butterfly taxonomy. Never ﾂ are herbivores, body smooth without hardened spines, theless, the "true" butterflies (superfamily Papili0- and possess extrusible glands (osmeteria) that are noidea) may be placed conservatively into four families unique among butterflies; pupa typically with a silk (Papilionidae,Pieridae, Nymphalidae, and Lycaenidae) girdle at third thoracic segment. that together include between 12,900 and 15,819 spe ﾂ Subfamily Baroniinae: one species (Baronia brevi- cies. One useful framework for organizing butterflytax- comis) endemic to Mexico; caterpillarsunique among 0nomic diversity is P. R. Ackery's (1984) synthesis of the family by feeding on Fabaceae. butterfly classification, which forms the basis of the Subfamily Parnassinae: 40-50 species, mainly in followingsynopsis: north temperate mountains; host plant families include Crassulaceae, Fumariaceae,and Zygophyllaceae. Family Papilionidae (Swallowtails, Fig. I): a group Subfamily Papihoninae (Fig. I): more than 500 of 500-600 species in three subfamilies, distributed species placed into three or more tribes; distributed worldwidebut with most species being tropical;adults worldwide; most species are tropica1.Many species mediumto large sized, both sexes have six walking legs with long hindwing tails; many with sexes strongly bearing nonbifid tarsalclaws; most are brightly colored, dimorphic; many involved in mimicry; species range may be Batesian or Mullerian mimics, all feed on flower from palatable mimics (Papilioand Eurytides) to unpa1- FIGURE1 Adult butterflies. Clockwise from upper left: (a) newly eclosed male Eurytldes protesi- 1aus (Papihomdae) drinking from wet soil; (b) male TherKas nr. hemon(Lycaenidae: Theclinae) perching on a leaf, (c) male Dismorphia amphiona(Piendae: Dismorphinae) visiting flowers; (d) female Mechanitis isthmia (Nymphalidae:Ithomnnae) ovipositing a cluster of eggs. [All photos copyright P. J. DeVnes.] . BUTTE RFLIES 561 atable models (Parides, Pachliopta, and Troides), and smal1,both sexes havesix walking legs (except Riodini- some are among the largest butterflies (.Trogonoptera nae), and frequently with alternating black and white and Omithoptera); host plant families include Annona- bands on the antennae; the group displays a tremendous ceae, Apiaceae, Aristolochiaceae, Fumariaceae,Her- diversity of form, color, and life histories; adults feed nandicaeae, Lauraceae, Magnoliaceae, Rosaceae,and on flower nectar, fruits, carrion, and honeydew, and a Rutaceae. few species do not feed as adults; some African and FamilyPieridae (whites, sulphurs, Jezebels, cabbage Southeast Asian groups are involved in mimicry com ﾂ butterflies;Fig. I): a group of more than 1100 species plexes; caterpillars most often slug-1ike, without hard in four subfamilies; adults small to medium-sized,both spines or projections, but the Riodininae shows an ex ﾂ sexes have six walking legs and distinctly bifid claws; tensive variety of form; as a group, lycaenidshave the most species are white or yellow or orange (or combina ﾂ widest･ diet breadth of all butterflies, and depending on tions thereof) derived from pterin pigments, some with the group caterpillarsmay feed on plants,other insects, red and black patterning; adults feed on flower nectar 0r insectsecretions; many caterpillarsform intimate and (Fig. 1), and males visit wet soil; caterpillars are herbi- complex symbiotic associations with ants and produce vores, body smooth without hardened spines, oftencov ﾂ acoustical calls similar to ant calls, secretions that are ered in granulations; pupa suspended at 45 ｡ angle from harvested by ants, and chemicals that alter ant behav- substratewith silk girdle at the first abdominal segment. iors;pupae are typicallyround, seed-shaped, unadorned Subfamily Pseudopontinae:one species (Pseudopon- with projections, and may produce clicking or whirring tia paradoxd) endemicto West Africa; early stage biolﾂ sounds when stimulated. ogy unknown. Subfamily Riodininae (metalmarks): 1200-1400 Subfamily Dismorphiinae: approximately 100 spe- species that are almost exclusively Neotropical; argu ﾂ cies, nearly all Neotropica1. Most Dismorphia (Fig. 1) ably the most diverse of all lycaenoid groups with re ﾂ are astonishingly precise mimics of certain aposematic spect to adult and larval forms, and they are often Nymphalidae(HeHconius, Mechanitis, and Oleria), and treated as a distinct family (Riodinidae) divided into represent the original example from which Batesian five subfamilies; adults feed primarily on flower nectar, mimicry theory was derived; host plants are in the some drink at wet soi1, others at carrion; overall their Fabaceae. life histories are poorly known, but as a group riodinid Subfamily Pierinae(whites, Jezebe1, and cabbage but- caterpillarsappear to be mainly herbivores(Euselasia, terflies): approximately 700 mostly tropicalspecies, es ﾂ Me5 山cmM ， An り @M5 ， M打 aCh& 小 ， Mぬ 伽 e， り川 mdChM ， pecially in Africa and Asia; some species have spectacu ﾂ Emesis,Anteros, and Helicopis), with a few carnivorous larly bright, contrasting colors of white, red, and black species on Homoptera (Alesa and Setabis); about (e.g., Delias, Mylothris, Pereute, and allies) suggesting one-third of the riodinids form symbioses with ants unpalatability and warning coloration; some groups and have vibratory papillae as sound producing organs with dimorphic sexes in which females resemble unpal ﾂ [Thisbe,Audre, Lemonias, Synargis, Nymphidium, and atable nymphalids and papilionids; some species un- 5etabis (Fig. 2)]; riodinid myrmecophilyis entirely dergo spectacular mass migrations (e.g., Ascia, Appias, Neotropical; host plant families include Araceae, and Belenois), and a few species are crop pests (Pieris); Asteraceae, Bromeliaceae, Bombacaceae, Cecropiaceae, host plant families include Capparidaceae, Brassicaceae, Clusiaceae, Dilleniaceae, Euphorbiaceae, Fabaceae, Santalaceae, and Loranthaceae. Lecythidaceae, Loranthaceae, Malpighiaceae, Maranta- Subfamily Coliadinae (sulphurs): 400-600 species ceae, Melastomataceae, Myrtaceae, Orchidaceae, Rubia- well represented in temperate and subtropical regions; ceae, Sapindaceae, and Zingiberaceae plus mosses, generally yellow or white with short, thickly scaled 1iverworts, and lichens. antennae; common in open areas, some migrate in large Subfamily Styginae: one enigmatic species (.Styxin- numbers and have been recorded out at sea far from femalis) that is endemic to the Peruvian high Andes; land (Colias, Phoebis, Catopsilia, Aphrissa, and Euremd); the early stage biology is unknown. many species with patterns visible only in the ultra ﾂ Subfamily Lipteninae: a small African group of violet. 30-40 species; some are Batesian mimics of aposematic Family Lycaenidae (hairstreaks, blues,