Dione Vanillae
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The Gulf fritillary or passion butterfly (''Dione vanillae'') is a bright orange
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
in the subfamily
Heliconiinae The Heliconiinae, commonly called heliconians or longwings, are a subfamily of the brush-footed butterflies (family Nymphalidae). They can be divided into 45–50 genera and were sometimes treated as a separate family Heliconiidae within the Pa ...
of the family
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a red ...
. That subfamily was formerly set apart as a separate family, the
Heliconiidae The Heliconiinae, commonly called heliconians or longwings, are a subfamily of the brush-footed butterflies (family Nymphalidae). They can be divided into 45–50 genera and were sometimes treated as a separate family Heliconiidae within the ...
. The Heliconiinae are "longwing butterflies", which have long, narrow wings compared to other butterflies. ''Dione vanillae'' is most commonly found in the southern areas of the United States, specifically in many regions of
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
and
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
. Gulf fritillaries have a
chemical defense Chemical defense is a life history strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites or chemical warnings which incite defensive behavioral changes. The production of defensive chemicals occurs in ...
mechanism in which they release odorous chemicals in response to predator sightings. As a result, common predators learn to avoid this species.
Pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
play a critical role in male-female courtship behaviors, with male gulf fritillaries emitting sex pheromones that contribute to mate choice in females. The scientific name ''Papilio vanillae'' was given to the gulf fritillary by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in 1758, based on a 1705 painting by
Maria Sibylla Merian Maria Sibylla Merian (2 April 164713 January 1717) was a German naturalist and scientific illustrator. She was one of the earliest European naturalists to observe insects directly. Merian was a descendant of the Frankfurt branch of the Swiss Mer ...
(''The Metamorphosis of the Insects of Surinam'', Plate XXV), which shows the adult and caterpillar of the gulf fritillary on a vanilla orchid, ''
Vanilla planifolia ''Vanilla planifolia'' is a species of vanilla orchid. It is native to Mexico and Belize. It is one of the primary sources for vanilla flavouring, due to its high vanillin content. Common names include flat-leaved vanilla, and West Indian vanil ...
''. However, It is now known that the gulf fritillary caterpillar does not use the vanilla plant.Shapiro, Arthur M., Journal of the Lepidopterists’ Society 62(1), 2008, 58–59. The species was moved to the genus ''Agraulis'' in 1835 by
Boisduval Jean Baptiste Alphonse Déchauffour de Boisduval (24 June 1799 – 30 December 1879) was a French lepidopterist, botanist, and physician. He was one of the most celebrated lepidopterists of France, and was the co-founder of the Société entomol ...
& Le Conte. A phylogenetic analysis reported in 2019 placed ''Agraulis'' as a sub-genus of ''Dione''.


Description

In its adult form, the gulf fritillary is a medium-sized butterfly that has extended
forewings Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwin ...
and a
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan of ...
range of 6.5 to 9.5 cm. This butterfly exhibits
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
as females are typically distinctively larger in size than males. The underside of the wings is brown and speckled with silvery white dots. In contrast, the top surface layer of the wings is deep orange in color with black streaks running across. In addition to the size difference between the two sexes, females are usually darker in color and are more marked with black streaks as compared to the males.


Geographic range

Gulf fritillaries are found primarily in the southern parts of the
U.S. The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
, such as many regions of
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
and
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. However, this butterfly’s range can extend from the southern U.S. into parts of
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
and sometimes as far as parts of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. They are also found in Hawaii. They have become increasingly common in California’s Central Valley since 2010.


Habitat

Gulf fritillaries are found in open
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s, like in moderately sunny areas near open grasslands, parks, and woodlands. Gulf fritillaries are also commonly seen in local butterfly gardens across the U.S.


Host plants

The caterpillar food plants--also called the host plants--for gulf fritillaries are members of the genus ''
Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...
''. The adult butterflies use nectar from many flowers, including ''
Lantana ''Lantana'' () is a genus of about 150 species of perennial flowering plants in the verbena family, Verbenaceae. They are native to tropical regions of the Americas and Africa but exist as an introduced species in numerous areas, especially in ...
'' plants. The Passiflora host plants are frequently called passion vines; in some Texan counties where this butterfly's population is high, gulf fritillaries will feed on specific species such as ''
Passiflora lutea ''Passiflora lutea'', commonly known as yellow passionflower, is a flowering perennial vine in the family Passifloraceae, native to the central and eastern United States. The vine has three-lobed leaves and small, yellowish-green, fringed flowers ...
'' and ''
Passiflora affinis ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...
''. These passion vine plants are suitable host plants as they provide a good structure for larval host habitats which enables young populations of gulf fritillaries to be sufficiently nurtured and protected. The role of host plants is also integral to the
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 typical ...
of gulf fritillaries, as the female butterflies lay their
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
s on or near the host plant.


Propagation


Oviposition

The process of oviposition in gulf fritillaries begins with the female butterfly flying low and slightly above the vegetation (most likely the typical host plants like ''
Passiflora incarnata ''Passiflora incarnata'', commonly known as maypop, purple passionflower, true passionflower, wild apricot, and wild passion vine, is a fast-growing perennial vine with climbing or trailing stems. A member of the passionflower genus ''Passiflora ...
'' or ''
Passiflora lutea ''Passiflora lutea'', commonly known as yellow passionflower, is a flowering perennial vine in the family Passifloraceae, native to the central and eastern United States. The vine has three-lobed leaves and small, yellowish-green, fringed flowers ...
'') and lasts until the female pauses above an individual plant. Then, the female makes contact with the host plant by landing; the female then deposits an egg to complete the process of oviposition. The driving factor behind what causes the female to oviposit on or near the host plant is most likely due to the certain chemical composition of the specific
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of the host plant. Once the female recognizes the chemical composition (by using the antennae), the female will oviposit.


Life cycle


Egg

Eggs are small and spherically shaped; the female gulf fritillary lays the eggs individually one by one on or near the host plant. Typical host plants include several species of the genus ''
Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...
''. The eggs are initially yellow in color after deposition by the female, and they gradually turn a brownish red color over the course of the next 24 hours. The average duration of the egg stage in the
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring *Life-cycle hypothesis, ...
for gulf fritillaries is roughly between three and five days depending on the temperature of the environment. Eggs are usually found on the upper surfaces of the leaves of the host plant, but some can be found on the underside of leaves as well.


Larva

In appearance, the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
e are dark orange with small black spines protruding outward from the body. The larval stages of gulf fritillaries include five
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
s, each with a varying timeframe. After the eggs hatch, the new larvae eat the egg casings for food. Eventually, the larvae will transition either to consuming other eggs or parts of the host plant for nourishment. Throughout the first three instar stages, the larvae primarily consume the
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
and feed away from the leaf margins of the host plant. How much of the leaf and how deep into the layers the larva will eat depend on the type (i.e. thickness or thinness) of the leaf in question. Throughout the last two instar stages, the larvae mainly consume the leaves and feed at the leaf margins. Throughout this food acquisition process, the degree to which a specific leaf is eaten before it is abandoned in favor of another newer leaf largely depends on whether food is in plentiful or scarce supply. The average duration of the larval stage is between 11 and 16 days depending on the temperature.


Pupa

In form and appearance, the
pupa A pupa ( la, pupa, "doll"; plural: ''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 ...
e of gulf fritillaries are not recognizably uniform, but rather can vary in both coloring and physical size. The average duration of the pupal stage in gulf fritillaries is roughly between 7 and 12 days depending on the temperature of the environment. Colder temperatures are associated with a longer time frame whereas warmer temperatures are associated with a shorter time for pupal development.


Metamorphosis

When the time comes for the caterpillar to create its
chrysalis A pupa ( la, pupa, "doll"; plural: ''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 ...
it turns a grayish color and begins to spin a silk-like substance into a ball on top or against a malleable surface. It then attaches its rear end to the "silk" lump and hangs upside down in a "j" position. By small contractions of the muscles it begins to shed its skin and head revealing a soft pinkish tan form. Quickly the soft form hardens and becomes grayish brown. The chrysalis stays in this form for eleven to twenty-one days. After that period of time, a small crack begins to form at the tip of the chrysalis revealing the butterfly's head. It continues to slowly move down through the bottom of the chrysalis until its legs are free to cling onto the shell of the chrysalis and pull itself the rest of the way out. Much like the monarch butterfly, it begins to pump the fluids from its bulging abdomen into its shriveled wings. When its wings are fully expanded it releases excess fluids from its abdomen. For the next ten to fifteen minutes it stays still and allows its wings to dry. Finally it fans its wings out and takes flight.


Adult

After the process of emerging from the
chrysalis A pupa ( la, pupa, "doll"; plural: ''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 ...
is completed, the butterfly dries its wings and eventually flies for the first time. In the adult population, the approximate
sex ratio The sex ratio (or gender ratio) is usually defined as 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. Many species devia ...
between males to females is roughly one to one, but further observational studies of gulf fritillaries are needed to better support the hypothesis of an equal sex ratio between the two dimorphic sexes. The average duration of the adult stage is roughly 14 to 27 days after the butterfly emerges from the chrysalis depending on the quality of the environment (i.e. factor in the availability of food, the presence of predators, competition between individuals).


Migration

With
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
constituting a major component of the geographic range of this butterfly, the
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
pattern of gulf fritillaries in Florida is well-documented. In Florida, gulf fritillaries have two major flights during the year. The first major migration involves huge populations of butterflies flying
north North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography. Etymology T ...
ward while the second migration involves the butterflies moving
south South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
ward throughout the state, especially throughout the
peninsula A peninsula (; ) is a landform that extends from a mainland and is surrounded by water on most, but not all of its borders. A peninsula is also sometimes defined as a piece of land bordered by water on three of its sides. Peninsulas exist on all ...
r parts of Florida. The first and second migrations occur in the spring and the fall, respectively.


Defense against predators

Both male and female gulf fritillaries possess certain defensive
gland In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland). Structure De ...
s located on their
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
s that serve as a
defense mechanism In psychoanalytic theory, a defence mechanism (American English: defense mechanism), is an unconscious psychological operation that functions to protect a person from anxiety-producing thoughts and feelings related to internal conflicts and o ...
against
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
, especially avian predators such as
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s. When the butterflies sense danger in the area or are suddenly disturbed, these glands emit a distinct and obvious
odor An odor (American English) or odour (English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is caused by one or more volatilized chemical compounds ...
which is composed of several different types of chemicals, such as 6-methyl-5-hepten-2-one and hexadecyl acetate. The summation of these chemical compounds released from the glands leads to birds preferentially avoiding the gulf fritillaries in favor of other prey. In some cases, gulf fritillaries can sometimes be subjected to
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
and
fighting Combat ( French for ''fight'') is a purposeful violent conflict meant to physically harm or kill the opposition. Combat may be armed (using weapons) or unarmed ( not using weapons). Combat is sometimes resorted to as a method of self-defense, or ...
from other butterfly species, such as ''Heliconius charithonia vazquezae'' and ''Dryas iulia moderata'' when those species have breeding populations in similar areas and within the same geographic range.


Mating


Courtship behavior

The general process for a typical courtship interaction begins when a male flies and lands near a perching female, who is most likely perched on a host plant, such as several species of the
Passiflora ''Passiflora'', known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae. They are mostly tendril-bearing vines, with some being shrubs or trees. They ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
. Once the male has landed, the male assumes a position next to the female with their heads together and with their bodies aligned at a 45-degree angle. At this time, the male engages in a specific action called the wing clap display in which the male continuously claps its wings open and closed. During this time, the
antenna Antenna ( antennas or antennae) may refer to: Science and engineering * Antenna (radio), also known as an aerial, a transducer designed to transmit or receive electromagnetic (e.g., TV or radio) waves * Antennae Galaxies, the name of two collid ...
e of the female are placed between the opening and closing wings. After the male ceases wing movement, the male butterfly will move into a position that facilitates genital contact with the female. The average duration of the copulation process including the wing clap display is approximately 11 seconds.


Female and male interactions


Pheromones

The release of
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s by the male gulf fritillaries plays a critical role in the mating and courtship process. Studies have shown that the males possess specific structures on their dorsal wings that engage in the production and eventual release of chemical pheromones. It is likely that pores that exist on the wings of the male butterflies transport the pheromones to the surface where the chemicals can be released and can then affect the females. The use of pheromones in the mating process relates to the idea of
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 Choic ...
.


Nuptial gifts

Along with chemical pheromones, the presence of
nuptial gift A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices. Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simpl ...
s plays an important role in the mating and courtship process. During the physical act of copulation, males transfer nutritious materials to the females. These extra nutrients given to the females can be used by a female to her advantage during the process of
oogenesis Oogenesis, ovogenesis, or oögenesis is the differentiation of the ovum (egg cell) into a cell competent to further develop when fertilized. It is developed from the primary oocyte by maturation. Oogenesis is initiated in the embryonic stage. O ...
. The use of nuptial gifts in the mating process, especially when females mate with multiple males, relates to the idea of
female 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 Choic ...
in deciding egg
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
.


Gallery

File:Gulf Fritillary Proboscis.jpg, Feeding using its proboscis File:Fritillary in sunlight3.jpg, Profile of wings in sunlight File:Gulf Fritillaries Mating 0019.jpg, Fritillaries mating File:Agraulis vanillae egg.jpg, Egg File:Agraulis vanillae (gulf fritillary) caterpillar.jpg, Caterpillar File:Agraulis vanillae vanillae, caterpillar.jpg File:Fritillary feeding on passion flower.jpg, Fritillary feeding on nectar from passion flower File:Agraulis vanillae chrysalis.jpg, ''Agraulis vanillae'' in its chrysalis form, seen in
Inglewood, California Inglewood is a city in southwestern Los Angeles County, California, in the Los Angeles metropolitan area. As of the 2020 U.S. Census, the city had a population of 107,762. It was incorporated on February 14, 1908. The city is in the South Bay ...
File:Agraulis vanillae larvae.jpg, ''Agraulis vanillae'' larvae preparing for their metamorphoses File:Gulf fritillary (Agraulis vanillae insularis).JPG, ''A. v. insularis''
Grand Cayman Grand Cayman is the largest of the three Cayman Islands and the location of the territory's capital, George Town. In relation to the other two Cayman Islands, it is approximately 75 miles (121 km) southwest of Little Cayman and 90 miles ( ...
File:Metamorphosis insectorum surinamensium (Pl. 25) BHL41398774.jpg, Plate XXV in Maria Sibylla Merian's ''Metamorphosis insectorum surinamensium'' File:D. vanillae.tif, Pinned ''D. vanillae'' from Central GA (scale bar: 1 cm)


References


External links


Tree of Life: ''Agraulis vanillae''


University of Florida IFAS

San Diego Natural History Museum
Gulf friillary
Butterflies of Canada
How to Raise Gulf Fritillary Butterflies
WikiHow {{Taxonbar, from=Q134532 Heliconiini Butterflies of North America Butterflies of Central America Butterflies of the Caribbean Nymphalidae of South America Butterflies of Cuba Lepidoptera of Argentina Lepidoptera of Brazil Lepidoptera of Colombia Lepidoptera of Ecuador Lepidoptera of Venezuela Insects of Puerto Rico Fauna of the Amazon Fauna of the Eastern United States Fauna of the Western United States Fauna of the California chaparral and woodlands Butterflies described in 1758 Taxa named by Carl Linnaeus