''Papilio polyxenes'', the (eastern) black swallowtail, American swallowtail or parsnip swallowtail, is a
butterfly
Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
found throughout much of North America. An extremely similar-appearing species, '' Papilio joanae'', occurs in the
Ozark Mountains
The Ozarks, also known as the Ozark Mountains, Ozark Highlands or Ozark Plateau, is a physiographic region in the U.S. states of Missouri, Arkansas, and Oklahoma, as well as a small area in the southeastern corner of Kansas. The Ozarks cover ...
region, but it appears to be closely related to ''
Papilio machaon
''Papilio machaon'', the Old World swallowtail, is a butterfly of the family Papilionidae. The butterfly is also known as the common yellow swallowtail or simply the swallowtail (a common name applied to all members of the family, but this speci ...
'', rather than ''P. polyxenes''. The species is named after the figure in
Greek mythology
Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
,
Polyxena
In Greek mythology, Polyxena (; ) was the youngest daughter of King Priam of Troy and his queen, Hecuba. She does not appear in Homer, but in several other classical authors, though the details of her story vary considerably. After the fall of ...
(pron.: /pəˈlɪksɨnə/; Greek: Πολυξένη), who was the youngest daughter of King
Priam
In Greek mythology, Priam (; , ) was the legendary and last king of Troy during the Trojan War. He was the son of Laomedon. His many children included notable characters such as Hector, Paris, and Cassandra.
Etymology
Most scholars take the e ...
of Troy. Its caterpillar is called the parsley worm because the caterpillar feeds on
parsley
Parsley, or garden parsley (''Petroselinum crispum''), is a species of flowering plant in the family Apiaceae that is native to Greece, Morocco and the former Yugoslavia. It has been introduced and naturalisation (biology), naturalized in Eur ...
.
The ''Papilio polyxenes'' demonstrates
polyandry
Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives ...
and a
lek mating
A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. It can also refer to a space us ...
system, showing no male parental care and display sites. Females are therefore able to choose males based on these sites and males are the only resource the females find at these sites.
Taxonomy
''Papilio polyxenes'' is part of the tribe Papilionini of the swallowtail butterfly family,
Papilionidae
Swallowtail butterflies are large, colorful Butterfly, butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includ ...
. The members of this tribe all have tails on the hindwings, and therefore include species named swallowtail."Genus Papilio." http://en.butterflycorner.net/Genus-PAPILIO.366.0.html#c4148 ''P. polyxenes'' is part of the genus ''
Papilio
''Papilio'' is a genus in the swallowtail butterfly family, Papilionidae, as well as the only representative of the tribe Papilionini. The word ''papilio'' is Latin for butterfly.
It includes the common yellow swallowtail ('' Papilio machaon'') ...
'', which is the biggest group of the family Papilionidae. Subspecies ''polyxenes'', once found in Cuba, is now considered likely extinct. Members of this genus typically feed on plants of the family
Lauraceae
Lauraceae, or the laurels, is a plant Family (biology), family that includes the bay laurel, true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genus (biology), genera worldwide. They are dicotyled ...
,
Rutaceae
The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE in Bo ...
and
Umbelliferae
Apiaceae () or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus '' Apium,'' and commonly known as the celery, carrot, or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering ...
.
Distribution
''Papilio polyxenes'' are found from southern Canada through to South America. In North America they are more common east of the Rocky Mountains. They are usually found in open areas like fields, parks, marshes or deserts, and they prefer tropical or temperate habitats.
Morphology
Eggs and larvae
Eggs are pale yellow. Young larvae are mostly black and white with a saddle, and older larvae are green with black transverse bands containing yellow spots.
Caterpillar and chrysalis
This caterpillar absorbs toxins from the host plants, and therefore tastes poor to bird predators. The black swallowtail caterpillar has an orange "forked gland", called the
osmeterium
The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
. When in danger, the osmeterium, which looks like a
snake
Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
's tongue, everts and releases a foul smell to repel predators.
The
pupa
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
e may be green or brown, but not depending on surroundings or the background on which they have pupated. The color of the
chrysalis
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages the ...
is determined by a local genetic balance that ensures the majority of pupae will blend in.Black Swallowtail Butterflies of Canada A section of the green pupae will turn a much darker green at the very end of the pupae stage. This color change occurs a few hours to a full day before hatching. Unusually, this butterfly's chrysalis is girdled with a loop, with its feet attached to the silk pad, which helps the butterfly to unfold its very large wings when crawling out.
Sexual dimorphism
The black swallowtail has a wingspan of 6.9–8.4 cm, and females are typically larger than males. The upper wing surface is black with two rows of yellow spots – these spots are large and bright in males and smaller and lighter in females. Females have a prominent blue area between these two rows, while males have a much less prominent blue area. These differences give rise to effective
Batesian mimicry
Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, who worked on butt ...
of ''
Battus philenor
''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many di ...
'' seen in females.
Both sexes show a red spot with a black bullseye on the inner hind margin of the hindwings and an isolated yellow spot on the front edge of the wings. The
ventral
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
side of wings of males and females are essentially identical: forewings have two rows of pale yellow spots, and hindwings have rows of bright orange spots separated by areas of powdery blue. The ventral side also acts as an effective mimic for both males and females for protection against predators.
Mimicry
Female markings are similar to those of '' B. philenor'', allowing females to engage in dorsal
mimicry
In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...
to reduce risk of predation by birds that preferentially prey on the black swallowtail. Females have evolved dorsal mimicry because they spend more time revealing their dorsal wing side during oviposition. The ventral wing surface of the black swallowtail also mimics that of ''B. philenor'', so both males and females are protected when their ventral wing surface is displayed.
Intrasexual selection
Male black swallowtails can sometimes mimic the female wing-back pattern, and therefore succeed in reduced predation as well. However, males of the typical coloration are more successful in intrasexual competition for mating territories compared to the males who mimic the female wing pattern. Females have no preference based on wing markings, and are equally likely to mate with a typical versus an alternative coloration. Therefore, male-male
intrasexual selection
Sexual selection is a mechanism of evolution in which members of one sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (intrasexual sele ...
is of greater importance than female
mate choice
Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior.Bateson, Paul Patrick Gordon. "Mate Choice." Mate Choi ...
in maintaining the classic male wing-back coloration and pattern.
Life cycle
Females lay single eggs on host plants, usually on the new foliage and occasionally on flowers. The eggs stage lasts 4–9 days, the larval stage 10–30 days, and the pupal stage 18 days. The duration of these stages may vary depending on temperature and the species of the host plants.Minno MC, Butler JF, Hall DW. (2005). ''Florida Butterfly Caterpillars and their Host Plants.'' University Press of Florida. Gainesville, Florida. 341 pp.
Feeny ''et al.'', 1985 finds that survival of eggs and larvae of ''P. polyxenes'' varies depending on the location (near
Ithaca, New York
Ithaca () is a city in and the county seat of Tompkins County, New York, United States. Situated on the southern shore of Cayuga Lake in the Finger Lakes region of New York (state), New York, Ithaca is the largest community in the Ithaca metrop ...
) where the eggs were laid. Eggs laid on
wild carrot
''Daucus carota'', whose common names include wild carrot, European wild carrot, bird's nest, bishop's lace, and Queen Anne's lace (North America), is a flowering plant in the family Apiaceae. It is native to temperate regions of the Old World ...
or
parsnip
The parsnip (''Pastinaca sativa'') is a root vegetable closely related to carrot and parsley, all belonging to the flowering plant family Apiaceae. It is a biennial plant usually grown as an annual. Its long taproot has cream-colored skin an ...
plants were more likely to survive to the fifth
instar
An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
than were eggs laid on
poison hemlock
''Conium maculatum'', commonly known as hemlock (British English) or poison hemlock (American English), is a highly poisonous flowering plant in the carrot family Apiaceae, native to Europe and North Africa. It is herbaceous, with no woody parts, ...
. Eggs laid on plants at elevations of above sea-level were more likely to survive than eggs laid at 120 meters above sea-level. There were also significant differences in survival rates between early and late broods in a year, and between years. In all cases, the primary cause of mortality was predation.
Emergence
Winter is spent in the chrysalis stage, and adults will emerge in the spring to seek out host plants. Adults will emerge in the mornings on a daily basis. First brood adults will fly from mid-May until late June, second brood adults will fly from early July until late August, and occasionally a partial third brood will occur that will emerge later in the season.
Life expectancy
Members of the black swallowtail are long lived compared to other butterflies that inhabit temperate zones. They encounter little predation and are quick and agile if they are disturbed. However, mortality from predators will occur during roosting and during unfavorable weather due to the associated increase in predation. Adult butterflies are at the highest risk for predation when they are incapable of flight or are starved from poor weather.
Food plants
''Papilio polyxenes'' use a variety of herbs in the carrot family (
Apiaceae
Apiaceae () or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus ''Apium,'' and commonly known as the celery, carrot, or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering p ...
), but will choose the food plants for their larvae based on visual and chemical variations. Host plant odor is one of the cues involved in the selection of landing sites for
oviposition
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
. The responses to these cues are innate, and feeding on a host plant as a larva does not increase the preference for that plant as an adult.
Species of host plants include:
* American angelica, ''
Angelica atropurpurea
''Angelica atropurpurea'', known commonly as purplestem angelica, great angelica, American angelica, high angelica, and masterwort,
* Hairy angelica, '' Angelica venenosa''
*
Dill
Dill (''Anethum graveolens'') is an annual herb in the celery family Apiaceae. It is native to North Africa, Iran, and the Arabian Peninsula; it is grown widely in Eurasia, where its leaves and seeds are used as a herb or spice for flavouring ...
Berula erecta
''Berula erecta'', known as lesser water-parsnip, cutleaf waterparsnip, or narrow-leaved water-parsnip, is a member of the carrot family. Growing to around tall, it is found in or by water. It is widespread across much of Europe, Asia, Australi ...
Daucus carota
''Daucus carota'', whose common names include wild carrot, European wild carrot, bird's nest, bishop's lace, and Queen Anne's lace (North America), is a flowering plant in the family Apiaceae. It is native to temperate regions of the Old World ...
Fennel
Fennel (''Foeniculum vulgare'') is a flowering plant species in the carrot family. It is a hardy, perennial herb with yellow flowers and feathery leaves. It is indigenous to the shores of the Mediterranean but has become widely naturalized ...
Lovage
Lovage ( ; ''Levisticum officinale'') is a perennial plant, the sole species in the genus ''Levisticum'' in the family Apiaceae, subfamily Apioideae. It has been long cultivated in Europe and the leaves are used as a herb, the roots as a vegeta ...
Magnolia acuminata
''Magnolia acuminata'', commonly called the cucumber tree (often spelled as a single word "cucumbertree"), cucumber magnolia or blue magnolia, is one of the largest magnolias, and one of the cold-hardiest. It is a large forest tree of the Easte ...
''
* Sweetbay magnolia, ''
Magnolia virginiana
''Magnolia virginiana'', most commonly known as sweetbay magnolia, or merely sweetbay (also laurel magnolia, swampbay, swamp magnolia, white bay, or beaver tree), is a member of the magnolia family, Magnoliaceae. It was the first magnolia to be ...
Parsnip
The parsnip (''Pastinaca sativa'') is a root vegetable closely related to carrot and parsley, all belonging to the flowering plant family Apiaceae. It is a biennial plant usually grown as an annual. Its long taproot has cream-colored skin an ...
, ''Pastinaca sativa''
*
Parsley
Parsley, or garden parsley (''Petroselinum crispum''), is a species of flowering plant in the family Apiaceae that is native to Greece, Morocco and the former Yugoslavia. It has been introduced and naturalisation (biology), naturalized in Eur ...
Ruta graveolens
''Ruta graveolens'', commonly known as rue, common rue or herb-of-grace, is a species of the genus ''Ruta'' grown as an ornamental plant and herb. It is native to the Mediterranean. It is grown throughout the world in gardens, especially for it ...
''
* Water parsnip, ''
Sium suave
''Sium suave'', the water parsnip or hemlock waterparsnip, is a Perennial plant, perennial wildflower in the family Apiaceae. It is native to many areas of both Asia and North America. The common name water parsnip is due to its similarity to par ...
Zanthoxylum americanum
''Zanthoxylum americanum'', the common prickly-ash, common pricklyash, common prickly ash or northern prickly-ash (also sometimes called toothache tree, yellow wood, or suterberry), is an aromatic shrub or small tree native to central and easter ...
thoracic
The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen.
In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main ...
temperatures of around 24 degrees Celsius are necessary for flight. Therefore, the black swallowtail will regulate thoracic temperatures by behaviorally changing their abdomen position, wing position, orientation to the sun, perching duration, and perching height. In lower temperatures, butterflies will raise their abdomens above flattened wings, and will perch relatively close to the ground. In higher temperatures, butterflies will lower their abdomens in the shade of their wings. Higher temperatures are also associated with shorter perch durations, greater flight durations and higher perch heights.
Territorial defense
Male butterflies secure territories to use in mate location and courtship. These territories contain no significant concentration of
nectar
Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
sources, larval host plants or night settling sites. Once secured, a male will maintain exclusive use of a territory 95% of the time. Males will aggressively chase other males who approach their territory, and then return to their territory. Success in defending a territory depends on the number of competitors and his previous success, but the size of the male is not a contributing factor. Males that emerge early in the
brood
Brood may refer to:
Nature
* Brood, a collective term for offspring
* Brooding, the incubation of bird eggs by their parents
* Bee brood, the young of a beehive
* Individual broods of North American periodical cicadas:
** Brood X, the largest br ...
are more likely to defend a female-preferred territory. These males will have early access to available territories, and will choose the ones that are most preferred by females. What makes a territory desirable by females remains unknown, and is only measured by the number of aggressive encounters between males and the overall mating frequency at these sites.
Male territories are generally of high relative elevation and topographic distinctness. This feature serves as an advantage to the lek mating system described later, as males will be concentrated in predictable locations and will be easy to encounter by females.
Aggression
In previous studies, nearly 80% of successful courtship flights were confined to a male's territory. Because a preferred territory site is crucial in mating success, males are extremely aggressive in maintaining their territory. Black swallowtails have a 4:1 male biased
sex ratio
A sex ratio is the ratio of males to females in a population. As explained by Fisher's principle, for evolutionary reasons this is typically about 1:1 in species which reproduce sexually. However, many species deviate from an even sex ratio, ei ...
, and a low female mating frequency which leads to intense male-male competition.
Mating systems
Protandry
The black swallowtail is protandrous, meaning males emerge before females. This emergence pattern is advantageous, because males that emerge earlier have a greater success in competing for superior territories, indicated by female preference. These superior territories will most likely still be available for early emerging males, and securing one of these territories is highly predictive of mating success. Furthermore, female fertility is directly correlated with their weight at emergence. This favors larger females, and explains why they emerge later to prolong the larval feeding period. Male success is not dependent on size, so selection favors early emergence to get the best territories preferred by females, though this will most likely result in smaller males. However, there is a drawback to this emergence system. For biological reasons, overall male mating frequency decreases as the mating season goes on. Therefore, early emerging males with early access to preferred territories will not be able to mate as often later in the mating season when female emergence is at its peak.
Lek mating
This type of territorial organization leads the black swallowtail to engage in a lek mating system. These butterflies satisfy the four criteria for lekking behavior, as defined by J.W. Bradbury: (1) there is no male parental care, (2) males aggregate at specific sites for display, (3) the only resource females find at the lek are the males themselves and (4) females can select their mates.
The territory that has the most male-male encounters can be seen as being the most desirable to both males and females, and is also the territory that has the highest female visitation rate. Hilltop leks give the advantage to females because they make it easy to locate mates, and competition for superiority creates an array of males who have already demonstrated their quality as a mate.
Copulation
Males can only mate twice a day, but females will mate more than once to replace a sperm supply that has deteriorated with time. ''P. polyxenes'' has a long mating period due to females tendency to mating multiple times and having a broad emergence period. This allows males to mate several times during their lifetime, despite only being able to
copulate
Sexual intercourse (also coitus or copulation) is a sexual activity typically involving the insertion of the erect male penis inside the female vagina and followed by thrusting motions for sexual pleasure, reproduction, or both.Sexual inte ...
twice on the same day. The black swallowtail engages in brief courtship flights, and copulations will last around 45 minutes.
Similar species
* Old World swallowtail (''
Papilio machaon
''Papilio machaon'', the Old World swallowtail, is a butterfly of the family Papilionidae. The butterfly is also known as the common yellow swallowtail or simply the swallowtail (a common name applied to all members of the family, but this speci ...
The Papilio polyxenes was named the official state butterfly of
New Jersey
New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
on January 11, 2016."N.J. State Symbols." https://www.njstatehousetours.org/160/State-Symbols.
Gallery
File:Egglaying.JPG, Laying eggs on parsley
File:Swallowtaileggs.JPG, Eggs on parsley
File:Black Swallowtail larva.jpg, First-
instar
An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
caterpillar
Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths).
As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
in northeast
Georgia
Georgia most commonly refers to:
* Georgia (country), a country in the South Caucasus
* Georgia (U.S. state), a state in the southeastern United States
Georgia may also refer to:
People and fictional characters
* Georgia (name), a list of pe ...
File:Papilio polyxenes caterpillars Fifth instar.jpg, These second- or third-instar larvae in northeast Georgia still have spikes.
File:Black Swallowtail Papilio polyxenes Closeup.jpg, Late-instar caterpillar (without spikes) soon ready to pupate
File:Black-swallowtail-osmeterium.JPG,
Osmeterium
The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
visible after slight provocation
File:BlackSwallowtailColorVariations.jpg, Caterpillar color variations
File:Black Swallowtail Papilio polyxenes Chrysalis.jpg, Greenish-brown
chrysalis
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages the ...
File:Papilionidae - Papilio polyxenes.jpg, Ventral view with chrysalis
File:Papilio Polyxenes (Black Swallowtail - TX).jpg, alt=Black Swallowtail, Black Swallowtail in North Houston, TX
References
Eastern Black Swallowtail Butterfly: Large format reference quality (free) photographs Cirrus Digital Imaging
*
*
*
*Ferris, CD. and Emmel, J.F. (1982). Discussion of ''Papilio coloro'' W.G. Wright (= ''Papilio rudkini'' F. and R. Chermock) and ''Papilio polyxenes'' Fabricius. ''Bulletin of the Allyn Museum'' 76: 13 pp.
''This article is adapted in part fro at the
USGS
The United States Geological Survey (USGS), founded as the Geological Survey, is an government agency, agency of the United States Department of the Interior, U.S. Department of the Interior whose work spans the disciplines of biology, geograp ...