Overview

Brief Summary

Introduction

Introduction:

The swallowtail butterfly family, Papilionidae, consists of at least 550 species, many of which are large and colorful and recognizable even to non-specialists. While the majority of swallowtail species are found in tropical latitudes, representatives from the family can be found on every continent except Antarctica, and can be common in both tropical and temperate habitats. Swallowtail butterfly diversity is greatest in East and Southeast Asia, a region where many natural butterfly habitats are under extreme threat of destruction due to human activity. Some swallowtails, particularly representatives from the genus Parnassius, may fly at very high elevations. The birdwing butterflies (Troidini: Troides) of Australasia are the largest butterflies in the world. Collins and Morris (1985) provide an overview of the patterns of swallowtail diversity around the world. 

 The name "swallowtail" refers to a tail-like extension on the edge of the hindwing that is found in many, though not all, papilionids. The function of this tail is not known, but genetic studies in some species of Papilio suggest the tail is a labile character whose expression is controlled by a single gene (Clarke and Sheppard 1960, Clarke et al. 1968). 

 Within the Papilionidae, many families of larval hostplants are utilized, although five families generally dominate the host records: Aristolochiaceae, Annonaceae, Lauraceae, Apiaceae, and Rutaceae. Notably, the swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae) are limited almost exclusively to feeding on Aristolochiaceae. It has been demonstrated in a number of Aristolochia-feeders that caterpillars are able to sequester aristolochic acids, causing both the larval and adult stages to be unpalatable to predators (eg. von Euw et al. 1968). 

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Characteristics

Characteristics:

Three unambiguous morphological characteristics unite all papilionids and distinguish them from the other butterfly families. 

 
     
  1. All swallowtail caterpillars possess a forked, eversible organ behind the head known as an osmeterium. The osmeterium secretes a foul-smelling terpene-based defensive compound (Eisner and Meinwald 1965). When molested, a caterpillar will evert its osmeterium and rear its head back in an attempt to dissuade its antagonist. The specific composition of osmeterial secretions may vary between swallowtail species. 

       

    Image copyright 2002 Felix A. H. Sperling

  2.  
  3. The second anal vein (vein 2A) on the adult forewing extends to the wing margin and does not converge with the first anal vein (vein 1A). In all other butterfly families veins 1A and 2A fuse, and 2A does not reach the wing margin.    

    Drawing modified from Miller 1987, copyright 2002 R. D. Reed

  4.   
  5. The cervical sclerites join beneath the neck.
  6.  

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Diversity

Diversity description:

There are about 600 species of swallowtail butterflies (Scriber, 1984; Collings and Morris, 1985). These are generally recognized in three subfamilies: Baroniinae (a single species, Baronia brevicornis), Parnassiinae (recognized as 54-76 species in 8 genera), and Papilioninae (550 species in 17 genera)

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Introduction

The swallowtail butterfly family, Papilionidae, consists of at least 550 species, many of which are large and colorful and recognizable even to non-specialists. While the majority of swallowtail species are found in tropical latitudes, representatives from the family can be found on every continent except Antarctica, and can be common in both tropical and temperate habitats. Swallowtail butterfly diversity is greatest in East and Southeast Asia, a region where many natural butterfly habitats are under extreme threat of destruction due to human activity. Some swallowtails, particularly representatives from the genus Parnassius, may fly at very high elevations. The birdwing butterflies (Troidini: Troides) of Australasia are the largest butterflies in the world. Collins and Morris (1985) provide an overview of the patterns of swallowtail diversity around the world. 

 The name "swallowtail" refers to a tail-like extension on the edge of the hindwing that is found in many, though not all, papilionids. The function of this tail is not known, but genetic studies in some species of Papilio suggest the tail is a labile character whose expression is controlled by a single gene (Clarke and Sheppard 1960, Clarke et al. 1968).  

 Within the Papilionidae, many families of larval hostplants are utilized, although five families generally dominate the host records: Aristolochiaceae, Annonaceae, Lauraceae, Apiaceae, and Rutaceae. Notably, the swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae) are limited almost exclusively to feeding on Aristolochiaceae. It has been demonstrated in a number of Aristolochia-feeders that caterpillars are able to sequester aristolochic acids, causing both the larval and adult stages to be unpalatable to predators (eg. von Euw et al. 1968). 

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Comprehensive Description

Papilionidae (Swallowtail Butterflies)
These are large-sized butterflies. They are often black-and-yellow striped, or they are black with rows of yellow, blue, or red dots. The hindwings often have tails. Some species, such as the Battus philenor (Pipevine Swallowtail), are distasteful and toxic, while other species of swallowtail mimic this butterfly. The caterpillars tend to be smooth and green with markings from other colors. They have horn-like structures that appear when the caterpillar is disturbed, emitting a noxious smell. The caterpillars feed on a wide variety of plants, including members of the Carrot family, Wafer Ash and Prickly Ash, Spicebush and Sassafras, pipevines, wild cherries, and other trees. The pupae are angular-shaped, green or brown, and slung from a stem with a silk girdle.

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Characteristics

Three unambiguous morphological characteristics unite all papilionids and distinguish them from the other butterfly families.

  1. All swallowtail caterpillars possess a forked, eversible organ behind the head known as an osmeterium. The osmeterium secretes a foul-smelling terpene-based defensive compound (Eisner and Meinwald 1965). When molested, a caterpillar will evert its osmeterium and rear its head back in an attempt to dissuade its antagonist. The specific composition of osmeterial secretions may vary between swallowtail species.

    Image copyright © 2002 Felix A. H. Sperling

  2. The second anal vein (vein 2A) on the adult forewing extends to the wing margin and does not converge with the first anal vein (vein 1A). In all other butterfly families veins 1A and 2A fuse, and 2A does not reach the wing margin.

    Drawing modified from Miller 1987, copyright © 2002 R. D. Reed

  3. The cervical sclerites join beneath the neck.

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Distribution

Geographic Range

There are over 550 species of swallowtails around the world. Most are tropical, only about 30 are found in the United States, and only 8 are in Michigan.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native )

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Geographical Distribution

Geographic Range:

Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian

Geographic Range description:

Swallowtails have a worldwide distribution (Scriber, 1984; Collins and Morris 1985)

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Physical Description

Morphology

Physical Description

This family contains species that are the largest butterflies in North America. Swallowtails get their name from the "tails" on the back edge of their wings that reminded people of the forked tails of Swallows. Adults swallowtails have black or black and yellow/white wings, sometimes with additional blue or red markings. All Michigan species have the tails on their hindwings, but some species in other parts of North America don't have them.

Swallowtail caterpillars are large and smooth, but they have an orange "horn" that is hidden under the skin of the thorax. They pop it up when they are threatened by predators, and it gives off repellent chemicals. Young caterpillars are black with white spots, and look like bird droppings. As they get older they change color, then they are often green but with black or yellow stripes, and several species have large spots that look like eyes.

Some species of swallowtails are mimics of others that have toxic compounds in their bodies.

Other Physical Features: ectothermic ; bilateral symmetry

Sexual Dimorphism: female larger; sexes shaped differently

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Egg morphology

Texture:

smooth

Orientation:

upright

Egg mass pattern:

Usually laid singly on the foodplant, but occasionally in small clusters. Sometimes they are deposited near the foodplant rather than on it. (Scoble 1992)

Description of egg morphology:

Unsculptured and almost spherical.

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Larvae Morphology

Secondary setae:

present

Larval body description:

The body is sometimes smooth, but numerous secondary setae typically occur on the body (Scoble, 1992)

Crochet arrangement description:

The crochets are triordinal and, on the ventral prolets, arranged in a mesoseries. A weak, biordinal lateroseries may also occur on the ventral prolegs (Scoble, 1992)

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Pupa/Cocoon morphology

Pupa description:

Typically exposed and attached to its substrate by a silken girdle traversing the thorax.

Cremaster:

present

Cocoon:

absent

Cocoon description:

In Parnassiinae, pupation occurs in a loose cocoon on the ground (Munroe, 1982)

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Adult Thorax Morphology

Epiphysis:

present

Forelegs:

normal

Number of tibial spurs foreleg:

from 0

Number of tibial spurs midleg:

from 0 to 2

Number of tibial spurs hindleg:

from 2

Leg description:

foreleg is not reduced apart from the arolium and pulvilli

Forewing anal vein notation:

There are 2 anal veins that are united at the base of the wing but then diverge

Hindwing anal vein notation:

There is one anal vein

Forewing description:

From Scoble, 1992: "In the forewing R4 and R5 are typically stalked, CuP is usually present near the wing base.

Hindwing description:

From Scoble, 1992: "A humeral vein is present on the hindwing. Sometimes scent brushes occur in an anal fold of the male hindwing."

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Adult Head Morphology

Eyes:

smooth

Maxillary palpus:

minute

Female antennae:

clubbed

Male antennae:

clubbed

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Diagnostic Description

Synapomorphies

Apomorphies:

All synapomorphies below are directly from Scoble 1992.  A protrusible, forked osmeterium occurs on the larval prothorax.  In the adult, the arolium and pulvilli of the pretarsus are reduced.  In the forewing, vein 2A extends free to the margin whereas in other Papilionoidea and most other Lepidoptera, it fuses with 1A after a short distance, and fails to reach the wing margin.  The mesothoracic aorta lacks a horizontal chamber (unlike the condition found in Hesperioidea, Hedyloidea, and other Papilionoidea) but has ostiae (Hessel 1966).  The laterocervicalia are joined ventromedially, a feature absent from other butterflies (Ehrlich 1958; Miller 1987)

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Ecology

Habitat

Adult swallowtails fly in open areas (fields, vacant lots, meadows, open forest, sides of streams) near their food plants, especially where there are abundant flowers.

Caterpillars are found on their food plants.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: taiga ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: swamp

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Trophic Strategy

Food Habits

Swallowtail catepillars eat leaves and flowers of particular plans. One Michigan species only eats plants in the genus Aristolochia (Virginia Snakeroot, Dutchman's Pipe). Another prefers trembling aspen, another wild cherry. Several species eat plants in the carrot family.

Adult swallowtails sip nectar, but also mud and sometimes manure.

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Associations

Predation

Many of these species defend themselves with toxic chemicals, or mimic species that do. The caterpillars all produce chemicals the repel ants. They also are camouflaged, especially when they are young. The eyespots may startle or confuse a predator.

Adult swallowtails are strong fliers, and hide in trees at night.

Known Predators:

  • Aves
  • Soricidae (eat pupae)
  • Sigmodontinae (eat pupae)
  • Anura (eat adults)
  • Araneae, especially crab spiders and orb-weavers (eat adults)
  • Formicidae (eat caterpillars)
  • Hymenoptera (eat caterpillars and adults)
  • mantids (eat adults)
  • Diptera (eat caterpillars)
  • Coccinellidae (eat eggs)
  • Chrysopidae (eat eggs)
  • Acari (eat eggs)

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Life History and Behavior

Behavior

Communication and Perception

Like all butterflies, they mainly use scent and sight for communication. Males attract females with pheromones and special display flights. Females leave scent marks on plants where they've laid eggs, telling other female not to put their eggs there.

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Adult Behavior

Adult behavior:

diurnal

Adult behavior:

From Scoble 1992: "In many species the adults are sexually dimorphic. Usually the female is mimetic, and may be polymorphic with each morph mimicking a different model."

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Life Cycle

Development

Like all Moths and Butterflies, this family has complete metamorphosis. See More Information on Butterflies and Moths for an explanation of this. In cold climates they spend the winter as pupa in a chrysalis (no cocoon).

Development - Life Cycle: metamorphosis

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Life Expectancy

Lifespan/Longevity

These butterflies don't usually live more than a year. They often have two generations a year, with some adults emerging in the Spring, and then their offspring emerging as adults in Fall. This second generation only lives for a few months.

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Reproduction

After mating, females lay eggs one at a time on the underside of leaves of their food plants.

Breeding season: Spring to Fall

Key Reproductive Features: iteroparous ; seasonal breeding ; sexual ; fertilization (Internal ); oviparous

There is no parental care in this family.

Parental Investment: no parental involvement

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Life History: Immature Stages

Larval food habits description:

Five families generally dominate the host records: Aristolochiaceae, Annonaceae, Lauraceae, Apiaceae, and Rutaceae.

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Life History: Adults

Adult food habits description:

Feed at flowers while on the wing.

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Evolution and Systematics

Evolution

Systematic and taxonomic history

Systematic and taxonomic history:

Most authors accept three extant subfamilies: Baroniinae (containing just one species which occurs in Southwest Mexico); Parnassiinae (containing about 50 species, mostly in the Holarctic); and Papilioninae (containing 500+ species, mostly in old world tropics). Ehrlich (1958), Munroe (1961) and Hancock (1983) have discussed Papilionidae relationships.

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Fossil Record

Fossil record:

Durden and Rose (1978) propose a fourth subfamily of Papilionidae based on two species from middle Eocene deposits in Colorado, representing the extinct genus Praepapilio (Scoble, 1992).

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Discussion of Phylogenetic Relationships

View Papilionidae Tree

For several decades the favored view of Papilionidae has divided the family into four subfamilies: Baroniinae, Praepapilioninae, Parnassiinae, and Papilioninae (Munroe, 1961; Hancock, 1983), thought to be related (with constituent tribes) thusly:

Baroniinae and Praepapilioninae are both considered to be very primitive. The plesiomorphic nature of the sole species of the Baroniinae, Baronia brevicornis, has been supported by virtually all treatments. The sole genus of Praepapilioninae, Praepapilio, includes two species of extinct butterflies that were each described from single fossils found in a middle Eocene deposit in Colorado, U.S.A. (Durden and Rose, 1978). Praepapilio and Baronia share several striking similarities, but their exact relationship has been difficult to determine.

Both Praepapilio and Baronia have traditionally been considered to be the sister group to the rest of the swallowtails, but recently several alternative hypotheses regarding the root of the swallowtail tree have been considered. In an analysis of 103 morphological characters from 59 butterfly and 19 moth species, de Jong et al. (1996) presented some evidence that the genus Parnassius may be the sister group to the rest of the swallowtails. However, the intra-papilionid relationships in de Jong et al.'s (1996) study changed dramatically when the composition of the outgroups was altered, and a local analysis with a butterfly-only outgroup provided the traditional Baronia-as-sister rooting. Furthermore, de Jong et al. (1996) included only six papilionid species in the analysis, leaving out many taxa bearing characteristics key to understanding papilionid relationships.

Using a maximum-likelihood analysis of DNA sequence data, Caterino et al. (2001) found possible evidence for a root within the Papilioninae that would place Papilionini + Troidini as the sister group to the rest of the swallowtails. Because the likelihood of this intra-Papilioninae rooting was not found to be significantly higher than the traditional rooting, and because the traditional rooting is favored using maximum parsimony, we present the traditional rooting in our phylogeny.

Although a phylogenetic analysis of morphological characters has generally supported a monophyletic Parnassiinae (Miller, 1987), several other studies on morphological (Häuser, 1993; de Jong et al., 1996) and molecular (Yagi et al., 1999; Caterino et al., 2001) characters have called into question the monophyly of the subfamily. However, a new phylogenetic study on Parnassiinae, based on five mitochondrial and two nuclear genes, as well as 236 morphological characters (Nazari et al., 2007) has provided some evidence of a monophyletic Parnassiinae with three strongly supported tribes within the subfamily. This study also suggests that Praepapilio is sister only to Papilionini and Baronia is sister only to Parnassiinae, and likely none of the two are sister to all swallowtails as previously indicated.

With the exception of the intra-Papilioninae rooting alternative proposed by Caterino et al. (2001), the monophyly of the Papilioninae has been accepted by virtually every worker in the field. Despite this general consensus, the relationships of the tribes that comprise the subfamily have been debated. For our phylogeny, we follow the recommendation of Miller (1987) in recognizing Teinopalpini as the sister group to the Troidini + Papilionini group. At the time of this writing, no molecular data for Teinopalpini exist, and Miller (1987) provides the only rigorous phylogenetic analysis of the morphological data. The tribe Teinopalpini consists of only two species, both in the genus Teinopalpus. Munroe (1961) and Hancock (1983) did not grant Teinopalpini tribal status, but considered Teinopalpus to be a Graphiine genus (note that both authors used the name "Leptocircini" instead of "Graphiini").

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records:4347
Specimens with Sequences:3980
Specimens with Barcodes:3694
Species:606
Species With Barcodes:553
Public Records:2291
Public Species:406
Public BINs:303
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Barcode data

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Conservation

Conservation Status

One species in Michigan is considered threatened, it is the Pipe-vine Swallowtail, Battus philenor. It's existence is threatened in Michigan because its food plant has become rare. It is still common in other states. This species collects toxic chemicals from its food plant, and so birds won't eat it. Several other species are mimics of this species. If the toxic species disappears, the birds won't learn not to eat it, and the mimics won't get any protection from looking like it anymore.

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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Positive

These butterflies don't have strong effects on humans one way or the other. Sometimes they are a garden pest, eating carrots and related plants but this is not common. One species in the south is sometimes a minor pest of orange and lemon trees.

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Wikipedia

Swallowtail butterfly

This article is about the insect family. For other uses, see Swallowtail.
Swallowtail Butterfly

Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species.[1] Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the largest butterflies in the world, the birdwing butterflies of the genus Ornithoptera.[2]

Swallowtails have a number of distinctive features; for example, the Papilionid caterpillar bears a repugnatorial organ called the osmeterium on its prothorax. The osmeterium normally remains hidden, but when threatened, the larva turns it outward through a transverse dorsal groove[3] by inflating it with blood.

The forked appearance of the swallowtails' hind wings, which can be seen when the butterfly is resting with its wings spread, gave rise to the common name swallowtail. As for its formal name, Linnaeus chose Papilio for the type genus, as Papilio is Latin for 'butterfly'. For the specific epithets of the genus, Linnaeus applied the names of Greek heroes to the swallowtails. The type species: Papilio machaon honoured Machaon, one of the sons of Asclepius, mentioned in the Iliad.[4]

Distribution[edit]

As of 2005, 552 extant species have been identified,[1] which are distributed across the tropical and temperate regions of all continents except Antarctica. Various species inhabit altitudes ranging from sea level to high mountains, as in the case of most species of Parnassius. The majority of swallowtail species and the greatest diversity in form and lifestyle are found in the tropics and subtropical regions between 20°N and 20°S,[5]: particularly Southeast Asia, and between 20°N and 40°N in East Asia. Only 12 species are found in Europe[6] and only one species, Papilio machaon is found in the British Isles.[6] North America has 40 species, including several tropical species and Parnassius.[7]

The northernmost swallowtail is the Arctic Apollo (Parnassius arcticus), found in the Arctic Circle in northeastern Yakutia, at altitudes of 1500 meters above sea level.[8] In the Himalayas, various Apollo species such as Parnassius epaphus, have been found at altitudes of 6,000 meters above sea level.[9]:221

Appearance/Morphology[edit]

The detailed descriptions of morphological characteristics of the Papilionidae, as quoted in Bingham (1905) are as follows:[10]:1,2

Egg. "Dome-shaped, smooth or obscurely facetted, not as high as wide, somewhat leathery, opaque." (Doherty.)

Larva. Stout, smooth or with a series of fleshy tubercles on the dorsum : sometimes with a raised fleshy protuberance (the so-called hood or crest) on the fourth segment. The second segment has a transverse opening, out of which the larva protrudes at will and an erect, forked, glandular fleshy organ that emits a strong, penetrating, and somewhat pleasant odor.

Pupa. Variable in form but most often curved backwards. It is angulate, with the head truncate or rounded and the back of abdomen is smooth or tuberculate. It is attached by the tail, normally in a perpendicular position, and further secured by a silken girth round the middle. In Parnassius, the pupa is placed in a loose silken web between leaves.

Imago. Wings extraordinarily variable in shape. Hind wing very frequently has a tail, which may be slender, or broad and spatulate, but is always an extension of the termen at vein 4. In one genus, Armandia, the termen of the hind wing is prolonged into tails at the apices of veins 2 and 3 as well as at vein 4. Pore wing (except in the aberrant genera Parnassius and Hypermnestra) with all 12 veins present and in addition a short internal vein, vein 1 a,[11] that invariably terminates on the dorsal margin.

Stages of development of a papilionid – Giant Swallowtail (Papilio cresphontes)

Distinguishing characteristics[edit]

The key characteristics that differentiate the Papilionidae from the other butterfly families are:[2]

  • The presence of the osmeterium, a forked, fleshy eversible organ found in the prothoracic segment of papilionid caterpillars.
  • Venation in the adult forewing of papilionids is characteristic. In swallowtails, the second anal vein, 2A, extends up to the wing margin and does not link with the first anal vein, 1A. These veins are fused in other butterfly families and 2A does not reach the wing margin.
  • The sclerites of the cervix (membranous neck between the head and thorax) are fused beneath the neck where the muscles for head movement are anchored.

Special adaptations and defense[edit]

To defend against predators, swallowtail butterflies practice Batesian mimicry, a behavior in which the butterflies' appearance closely resemble that of distasteful species to ward off predators. Swallowtails differ from many animals that practice mimicry as certain species, such as the tiger swallowtail butterfly (Papilio glaucus), exhibit a female-limited polymorphism for Batesian mimicry and others, such as the Canadian tiger swallowtail (Papilio canadensis) do not display any form of mimicry.[12]

Predators of the swallowtail butterfly include the Red-winged blackbird, Pennsylvania firefly, Five-lined skink, Green darner, Goldenrod spider, Chinese mantid, Fiery Searcher, and Striped Skunk.[13]

Biological basis for polymorphisms in mimicry[edit]

In Papilio glaucus (Eastern Tiger swallowtail), Y-linkage determines whether the females are either wild-type (yellow and black) or melanic (dark melanin replaces the yellow background).[14] This genetic difference stems from the fact that melanism is controlled by a single gene, which controls the level of dopamine in the organism. During development, the enzyme BAS, which assists dopamine in producing the yellow pigmentation, normally found on the wings' background, is suppressed. Without the pigmentation, the butterfly appears mostly black (the melanic form) and is a Batesian mimic of Battus philenor, the Pipevine swallowtail. There are also Papilio glaucus that are not wholly black; several possess an intermediate "sooty" color and are sensitive to temperature.[12]

The different polymorphisms (wild-type, melanic, and the 'sooty' intermediate) depend upon the geographical distribution and abundance of its mimic, the Battus philenor, whose wing color varies depending on its geographical location.[12] In order to be successfully confused for the B. philenor by predators, the Papilio glaucus's background wing color matches that of the B. philenor residing in the same regional area. Studies support this theory; in the southeastern United States, the relative abundance of melanic females has been found to geographically correlate with B. philenor.

Influence of sexual harassment in wing polymorphism[edit]

Only certain subsets of swallowtail females practice mimicry. This polymorphism is seen in Papilo dardanus, the African swallowtail butterfly, whose females have three different morphs for wing color pattern: a black-and-white pattern for Batesian mimicry, a black and yellow pattern that resembles the males of the species, and a pattern with orange patches that resembles the elderly males of the species.[15] Given that the males of the species, which do not have Batesian mimicry, are preyed upon much more frequently by predators than the females, scientists initially questioned why females would choose an andromorph wing pattern, which would seemingly lower their fitness compared to the mimicry form.

Several hypotheses for this phenomena were made, the two noteworthy being the pseudosexual selection hypothesis and the male avoidance hypothesis. In the pseudosexual hypothesis, male butterflies aggressively approached the 'male' looking females and then mellowed their behavior into sexual behavior when they were close enough to identify them as females.[16] In the male avoidance hypothesis, female butterflies disguise themselves in an attempt to evade male harassment, as courtship can be harmful, time-consuming, and attract predators.[17]

One study recorded male responses to females of each morphs and found that the males consistently favored the Batesian mimics, then the black-and-yellow, and then the morph with orange patches.[15] The scientists concluded that frequency-dependent selection did lead to equal success for all three alternative strategies: the Batesian females suffered the least number of predators but their fitness was reduced the most by sexual harassment, while the other two faced lower sexual harassment but also lost fitness from predators' attacks.

Taxonomy[edit]

Subfamilies[edit]

The genera of extant swallowtails are usually classified into three subfamilies, Baroniinae, Parnassiinae, and Papilioninae, the latter two being further divided into tribes. In swallowtails, besides morphological characteristics, the choice of foodplants and ecological lifestyle reflect phylogeny and classification.

Baroniinae[edit]

The Baroniinae are a monotypic subfamily, restricted to a very small region in Mexico and are considered to be the most basal of the subfamilies. Baronia brevicornis is considered to be a relict species, and shares features with a fossil taxon Praepapilio. Baronia is unique amongst papilionids as having an Acacia species (family Leguminosae) as its foodplant.[5]

Parnassinae[edit]

The Parnassinae are a subfamily of essentially holarctic butterflies. The vast majority of species, mostly Parnassius, can be found in mountain habitats. Parnassinines can also be found in other habitats such as "arid deserts (Hypermnestra), humid forests (Luehdorfia) and even lowland meadows (Zerynthia)" (Nazari, 2006).[18] The tribes recognized in the Parnassinae are Parnassiini, Zerynthiini, and Luehdorfiini.

Tribe Parnassiini contains two monotypic genera, Hypermnestra, largely confined to central Asia and the genus Parnassius (the Apollos), a distinctive group of many species, all of which are alpine and capable of living at high altitudes. Most Parnassius have two small reddish spots on their hindwings. The tribe Luehdorfiini contains the monotypic genera Archon of Asia minor and the genus Luehdorfia of China and Japan. These two tribes have evolved to change their foodplants, while the third tribe, Zerynthiini, has retained the archetypical papilionid foodplant, the lowland vine Aristolochia. Zerynthhini comprises four genera – Sericinus, Bhutanitis, Zerynthia and Allancastria.[5]:13[19]

Subfamily : Parnassiinae.

Papilioninae[edit]

The tribes recognised in the Papilioninae are Leptocircini, Teinopalpini, Troidini, and Papilionini.

Subfamily : Papilioninae.

Praepapilioninae[edit]

An additional subfamily, Praepapilioninae, consisting of a single genus Praepapilio, includes two species of extinct butterflies, each member being described from single fossils found in a middle Eocene deposit in Colorado, U.S.A. (Durden and Rose, 1978).[20]

Phylogeny[edit]

A phylogeny of the Papilionidae based on Nazari (2007) is given:[2][19]

Phylogeny


Praepapilioninae (†)



Baroniinae



Parnassiinae

Parnassiini



Zerynthiini



Luehdorfiini



Papilioninae


Leptocircini



Teinopalpini





Papilionini



Troidini






Phylogeny of the Papilionidae
(after Nazari, 2007)[2][19]

It is now accepted that the subfamily Papilioninae is monophyletic.[2] The Swallowtail butterflies in the nominate tribe Papilionini number about 225 species and studies have been made on their host-plant coevolution and phylogeny. Old morphological classifications were also found to be valid in that they formed clusters. Species belonging to the groups that use Rutaceae as host plants formed two groups corresponding to Old World and American taxa. Those that fed on Lauraceae and Magnoliaceae were found to form another cluster which includes both Asian and American taxa.[21]

The Parnassinae, like the Papilioninae, were also believed to be monophyletic based on morphological studies but recent studies based on both morphological and molecular characteristics suggest that this is not the case.[2] Of the Parnassiinae, the genera Parnassius and Hypermnestra were found to be extremely close based on molecular studies[22] and are now considered to be part of the tribe Parnassiini.[19] The two taxa, Archon and Luehdorfia, have been found to be closely related through analysis of nuclear and mitochondrial DNA, and, though they share no morphological similarities, have now been united in the tribe Luehdorfiini.[19]

The subfamily Baroniinae is represented by the sole representative species Baronia brevicornis. They are unique in the family to use the Fabaceae (Leguminosae) as their larval host plants. The Baronninae and the extinct subfamily Praepapilioninae share many external similarities and are traditionally considered to be the most primitive families and sister to the rest of the swallowtails. Recent research suggests that this may not be the case, the Baroniinae being closely related to only the Parnassiinae, and Praepapilio to only the Papilionini and neither taxa being sister to the rest of the swallowtails.[2]

Mating and young[edit]

After mating, the male Parnassines produce a glue like substance that is used to seal the female genital opening and prevent other males from mating.[23] They lay individual eggs on the underside of the leaves of their food plants.[24] There is no parental investment once the eggs have been laid.

The pupae are typically attached to the substrate by the cremaster but with head up held by a silk girdle. The Apollos, however, pupate in debris on the ground and also build a loose cocoon. In the temperate regions, the winters are passed in a pupal diapause stage.

Food[edit]

Scarce Swallowtail butterfly, Iphiclides podalirius on lavender flowers, near Adriatic coast.

The caterpillars of various swallowtail butterfly species feed on a wide range of different plants, most depending on only one of five families: Aristolochiaceae, Annonaceae, Lauraceae, Umbelliferae (Apiaceae) and Rutaceae. By eating some of these toxic plants, the caterpillars sequester aristolochic acid which renders both the caterpillars and the butterflies of some of these as toxic, thus protecting them from predators.[25] Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae) almost exclusively use the Aristolochiaceae family as their host plants.

For example, the Eastern Black Swallowtail's main host plant in the wild is Queen Anne's Lace, but they also eat garden plants in the carrot family, including carrots, parsley, dill, and fennel.[13]

Adult swallowtails sip nectar, but also mud and sometimes manure.[24]

Swallowtails and humans[edit]

As swallowtail butterflies are large, colourful, and attractive, they were once targeteted by butterfly collectors. The largest of these, the Birdwing butterflies are particularly sought after and are cultured in butterfly farms for the purpose of collectors.

Many members of the family feed as larvae on plants of the citrus family, Rutaceae. Some of these attractive butterflies are therefore considered pests in citrus orchards.

The Oregon Swallowtail is the state insect of Oregon. The Eastern Tiger Swallowtail is the state insect of Virginia and the state butterfly of Georgia, Delaware, and South Carolina. The Black Swallowtail is the state butterfly of Oklahoma.

In popular culture[edit]

The American TV show Gilligan's Island had an episode[26] where an explorer[27] came to the island seeking a rare "pussycat swallowtail."

Swallowtails have appeared in various forms of Japanese entertainment, such as tokusatsu, manga and anime. In the 1996 Season of the popular Japanese tokusatsu Metal Hero Series B-Fighter Kabuto and the 1997 American show Beetleborgs Metallix, one of the B-fighters/Astral Borgs motifs was a swallowtail hence her Japanese designated name "B-Fighter Ageha".[28] In the manga and anime Bleach, Shinigami use Hell Butterflies to send messages and travel between Soul Society and the Living World (Earth); the same butterflies also guide souls during soul burials.[citation needed] These swallowtails are entirely black except for a few red markings on the wings, making them resemble Papilio protenor.[29] The butterfly-based entity Beautifly from the Pokémon anime series is pictured as a swallowtail. Swallowtail butterflies also appear in the manga xxxHolic which resemble yuuko ichihara.[citation needed] The antagonist Koushaku Chono (Papillon) in the manga/anime Busou Renkin features swallowtail-like imagery.

The band Rudolf Steiner, later known as Schwarz Stein, recorded a song entitled 黒揚羽 (lit. Black Swallowtail) on an early demotape. It is a track on the 2006 collaboration album Another Cell.[citation needed] The band Elvenking recorded a song titled "Swallowtail", which was released on their 2006 album, The Winter Wake.[citation needed] Paramore's 2009 release "Brand New Eyes" features a dissected swallowtail butterfly on its cover.

In the Japanese video game Bayonetta the main character Bayonetta is a witch who is in a pact with a demoness who takes on a Swallowtail form

References[edit]

  1. ^ a b Häuser, Christoph L.; de Jong, Rienk; Lamas, Gerardo; Robbins, Robert K.; Smith, Campbell; Vane-Wright, Richard I. (28 July 2005). "Papilionidae – revised GloBIS/GART species checklist (2nd draft)". Retrieved 8 November 2010. 
  2. ^ a b c d e f g Reed, Robert D.; Sperling, Felix A.H. (2006). "Papilionidae – The Swallowtail Butterflies". Tree of Life Web Project. Retrieved 7 November 2010. 
  3. ^ Richards, O. W.; Davies, R.G. (1977). Imms' General Textbook of Entomology: Volume 1: Structure, Physiology and Development Volume 2: Classification and Biology. Berlin: Springer. ISBN 0-412-61390-5. 
  4. ^ Salmon, Michael A., Marren, Peter, Harley, Basil. The Aurelian Legacy: British Butterflies and Their Collectors. page 252. Publisher: University of California Press. 2001. ISBN 978-0-520-22963-1
  5. ^ a b c Collins, N. Mark; Collins, Michael G. (1985). Threatened Swallowtails of the World: the IUCN red data book. IUCN Protected Area Programme Series. Gland, Switzerland and Cambridge, U.K.: IUCN. pp. 401 & 8 plates. ISBN 978-2-88032-603-6. Retrieved 22 October 2010. 
  6. ^ a b Coombs, Simon (30 September 2010). "European Butterfly checklist". http://www.butterfly-guide.co.uk/. Retrieved 11 November 2010. 
  7. ^ Brock, Jim P.; Kaufman, Kenn (2003). Butterflies of North America. Boston: Houghton Mifflin. ISBN 0-618-15312-8. 
  8. ^ Stumpe, Felix. "Parnassius arctica Eisner, 1968". Russian-Insects.com. Retrieved 9 November 2010. 
  9. ^ Mani, M. S. (1968). Ecology and Biogeography of High Altitude Insects. Volume 4 of Series entomologica. Springer. p. 530. ISBN 978-90-6193-114-0. Retrieved 9 November 2010. 
  10. ^ Bingham, C.T. (1905). The Fauna of British India including Ceylon and Burma – Butterflies (Vol 1). London: Taylor and Francis. p. 519. Retrieved 7 November 2010. 
  11. ^ The vein is since named 2A or second anal vein in modern venation systems.
  12. ^ a b c Scriber, Mark; Hagen, Robert; Lederhouse, Robert (February 1996). "Genetics of Mimicry in the Tiger Swallowtail Butterflies, Papilio glaucus and P. canadensis (Lepidoptera: Papilionidae)". Evolution 50 (1): 222. doi:10.2307/2410795. JSTOR 2410795. 
  13. ^ a b Moran, Mark. "Eastern Black Swallowtail". Retrieved 19 October 2013. 
  14. ^ Koch, Bernhardt; Behnecke, Bettina; ffrench-Constant, Richard H. (May 2000). "The molecuar basis of melanism and mimicry in a swallowtail butterfly". Current Biology 10 (10): 591–4. doi:10.1016/s0960-9822(00)00494-2. PMID 10837223. 
  15. ^ a b Cook, S. E.; Jennifer G. Vernon; Melissa Bateson; Tim Guilford (1994). "Mate Choice in the Polymorphic African Swallowtail Butterfly, Papilio dardanus: male-like females may avoid sexual harassment". Animal Behavior 47 (2): 389–397. doi:10.1006/anbe.1994.1053. Retrieved 15 September 2013. 
  16. ^ Vane-Wright, R.; C.R. Smith (1991). "Phylogenetic Relationships of Three African Swallowtail Butterflies, Papilio dardanus, P. phorcas, and P. constantinus: a cladistic analysis (Lepidoptera: Papilionidae)". Systematic Entomology. Biology of Butterflies 16 (3): 275–291. doi:10.1111/j.1365-3113.1991.tb00689.x. 
  17. ^ Conrad, K.F.; G Pritchard (1989). "Female Dimorphism and Physiological Colour Change in the Damselfly Argia vivida Hagen Odonata: Coenagrionidae)". Canadian Journal of Zoology 67 (2): 298–304. doi:10.1139/z89-044. 
  18. ^ Nazari, Vazrick (2006). "Parnassius Latreille 1804". Tree of Life Web Project. Retrieved 7 November 2010. 
  19. ^ a b c d e Nazari, Vazrick; Sperling, Felix A.H. (2006). "Parnassiinae Duponchel, [1835]". Tree of Life. Tree of Life Web Project. Retrieved 7 November 2010. 
  20. ^ Durden, C. J.; Rose, H. & Rothschild, Miriam (1978). "Butterflies from the middle Eocene: the earliest occurrence of fossil Papilionidae (Lepidoptera)". Pearce-Sellards Ser. Tex. Mem. Mus. 29 (5): 1–25 .
  21. ^ Aubert, J.; Legal, L; Descimon, H.; Michel, F. (1999). "Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)". Mol Phylogenet Evol. 12 (2): 156–167. doi:10.1006/mpev.1998.0605. PMID 10381318 .
  22. ^ Katoh, T.; Chichvarkhin, A.; Yagi, T.; Omoto, K. (2005). "Phylogeny and evolution of butterflies of the genus Parnassius: inferences from mitochondrial 16S and ND1 sequences". Zoolog Sci. 22 (3): 343–351. doi:10.2108/zsj.22.343. PMID 15795497 .
  23. ^ Ramel, Alain. "Les Papilionides, une famille en beauté". Les Insectes – Petit cours illustré d'entomologie(The Insects – A short illustrated course in Entomology). Retrieved 8 November 2010.  English translation.
  24. ^ a b "Swallowtail Butterflies". University of Michigan. Retrieved 19 October 2013. 
  25. ^ von Euw, J.; Reichstein, T. & Rothschild, M. (1968). "Aristolochic acid in the swallowtail butterfly Pachlioptera aristolochiae". Isr. J. Chem. 6: 659–670. doi:10.1002/ijch.196800084 .
  26. ^ "Gilligan's Island Script Episode #75, "Man With A Net"". episode #75 script. gilligansisle.com. 
  27. ^ "Gilligan's Island Season 3, Episode 7 summary". Man with a Net. tv.com. 
  28. ^ Though her insect designation was never announced in Beetleborgs Metallix, her name being Ladyborg, the astral coin that was used to summon her has the illustration of a swallowtail.
  29. ^ "Bleach Soul Reaper Guide". MyFavoriteGames.com. Retrieved 30 November 2009. 
  • Chattopadhyay, Jagannath, (2007),"Swallowtail Butterflies, Biology & Ecology of a few Indian Species, Desh Prashan, Kolkata, India",134pp.ISBN 978-81-905719-1-3.
  • Igarashi, S., 1979. Papilionidae and their early stages. Volume I Text (in Japanese), Volume 2 Plates. Kodansha, Tokyo.

Further reading[edit]

  • Glassberg, Jeffrey Butterflies through Binoculars, The West (2001)
  • Guppy, Crispin S. and Shepard, Jon H. Butterflies of British Columbia (2001)
  • James, David G. and Nunnallee, David Life Histories of Cascadia Butterflies (2011)
  • Pelham, Jonathan Catalogue of the Butterflies of the United States and Canada (2008)
  • Pyle, Robert Michael The Butterflies of Cascadia (2002)
  • Rothschild, 1895 A revision of the Papilios of the Eastern Hemisphere, exclusive of Africa Novit. Zool. 2 (3): 167-463 and plates pdf
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