Papilio Appalachiensis
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''Papilio appalachiensis'', the Appalachian tiger swallowtail, is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
swallowtail butterfly Swallowtail butterflies are large, colorful 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 includes the large ...
found in eastern North America, particularly in the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
. It is a hybrid of another two ''
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''), ...
'' species, ''
Papilio canadensis ''Papilio canadensis'', the Canadian tiger swallowtail, is a species of butterfly in the family Papilionidae. It was once classified as a subspecies of ''Papilio glaucus''. Description Adult The wingspan of this species is 67 to 80 mm. It ...
'' and ''
Papilio glaucus ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
'', with which it shares many characteristics. The butterflies are normally yellow and contain black patterns in their wings. Their wingspans range from 86 to 115 mm. The caterpillars range in color from green and yellow to orange and are ornamented with black specks that give them the appearance of a bird dropping, which is useful for camouflage, or a large eye, a form of mimicry that is also efficient for protection. This species is
univoltine Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...
. Females lay their eggs in May.


Taxonomy

''Papilio appalachiensis'' is a member of the
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 ...
''
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''), ...
'' in the order
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
. ''P. appalachiensis'' is considered to be a hybrid of two other members of the genus ''Papilio'': ''
Papilio canadensis ''Papilio canadensis'', the Canadian tiger swallowtail, is a species of butterfly in the family Papilionidae. It was once classified as a subspecies of ''Papilio glaucus''. Description Adult The wingspan of this species is 67 to 80 mm. It ...
'' and ''
Papilio glaucus ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
''.


Distribution

The butterfly is found in the eastern United States, specifically in the Appalachian Mountains, ranging from
Pennsylvania Pennsylvania (; ( Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, ...
to
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
. It is notably larger than both the
eastern tiger swallowtail ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
and the Canadian tiger swallowtail.


Evolution

''P. appalachiensis'' is thought to have evolved as a
hybrid species Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was tho ...
of two other ''
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''), ...
'' butterflies: '' P. canadensis'' and '' P. glaucus''. Originally, researchers believed that ''P. canadensis'' and ''P. glaucus'' were distributed in distinct regions separated by a
hybrid zone A hybrid zone exists where the ranges of two interbreeding species or diverged intraspecific lineages meet and cross-fertilize. Hybrid zones can form ''in situ'' due to the evolution of a new lineage but generally they result from secondary contact ...
stretching east from Minnesota to southern New England and south along the Appalachian Mountains. In 2005, researchers suggested these two species interacted at some point and produced the new hybrid ''P. appalachiensis'' in the hybrid zone. Although each species of tiger swallowtail butterflies is usually confined to a specific "thermal landscape", based on specific
X-linked Sex linked describes the sex-specific patterns of inheritance and presentation when a gene mutation (allele) is present on a sex chromosome (allosome) rather than a non-sex chromosome (autosome). In humans, these are termed X-linked recessive, ...
markers and various behavioral traits, laboratory and observational studies have shown the hybrid Appalachian tiger swallowtail butterflies have emerged. Data collected about
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 ...
preferences and larval mortality rates further suggest hybridization between ''P. canadensis'' and ''P. glaucus''. Due to shifting thermal landscapes, the ranges of both ''P. glaucus'' and ''P. canadensis'' have moved dramatically enough to overlay, leading to crossbreeding that has resulted in ''P. appalachiensis''. Although
hybrid speciation Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was th ...
is often difficult to achieve, given the difficulty of finding a habitat separate from either parental species (
competitive exclusion In ecology, the competitive exclusion principle, sometimes referred to as Gause's law, is a proposition that two species which compete for the same limited resource cannot coexist at constant population values. When one species has even the sligh ...
) and remaining genetically distinct from the parental species (
reproductive isolation The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring ...
), ''P. appalachiensis'' accomplishes this by living in a separate habitat: a higher elevation in the Appalachian Mountains.


Appearance

The Appalachian tiger swallowtail has similarities to both parent species; it resembles ''P. canadensis'' as it is adapted to a cooler temperature and has only one brood per year (
univoltine Voltinism is a term used in biology to indicate the number of broods or generations of an organism in a year. The term is most often applied to insects, and is particularly in use in sericulture, where silkworm varieties vary in their voltinism. ...
), but also resembles ''P. glaucus'' as it displays sex-limited polymorphism with the dark female morph displaying
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. Often, mimicry f ...
. However, research has shown that ''P. appalachiensis'' may be more closely related genetically to ''P. canadensis'' than to ''P. glaucus''. Females of ''P. appalachiensis'' are generally larger than those of ''P. glaucus''. Their wings are slightly angular and have vertical black stripes that are narrower than those on ''P. glaucus''. The hindwing scallops are also less curved and more angular than those of ''P. glaucus''. Additionally, the spots on their hindwings are also less crescent shaped and rectangular compared to those of ''P. glaucus''. Males' hindwings tend to be more triangular. The female has two forms: yellow and black. Both the yellow and black forms have a slight blue tint on the topside of the hindwings that is significantly reduced from the stronger blue tint on those of the tiger swallowtails. On the topside of the hindwing, there is a large orange spot. Their wingspan is . Initially, no black forms of ''P. appalachiensis'' were thought to exist, but researchers found two black females on the summit of
Spruce Knob Spruce Mountain, located in eastern West Virginia, is the highest ridge of the Allegheny Mountains. The whale-backed ridge extends for only from northeast to southwest, but several of its peaks exceed in elevation. The summit, Spruce Knob (4863 ...
in West Virginia.


Caterpillar

The
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 Sym ...
s of ''Papilio'' species are usually smooth and vary in color from green to yellow orange. In addition, they have black markings to mimic the appearance of bird droppings or the head of another bug with large "eyes". The eyespot consists of areas of yellow ringed with black with a smaller, light purple spot (also ringed in black) inside. Additionally, a small black spot on top of the "eye" gives it the appearance of a true eye. This form of
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. Often, mimicry f ...
provides a natural defense mechanism against potential predators. In addition to their appearance, every caterpillar of the swallowtail family has an
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 ...
, a special scent gland behind the head that, when disturbed, is everted from its pouch and releases a foul odor. The gland is orange red, fleshy, forked and generally intimidating to potential predators. The mimicry induced by the eyespots and forked gland give the caterpillar the appearance of a snake, adding to its defense mechanisms against potential predators. The caterpillar lives in a nest that it creates in the trees by folding a leaf over itself and pulling together the two halves with silk. By the time it is ready to undergo metamorphosis, the caterpillar can grow to be up to 5 cm (2 in) long. About a month is needed for the egg to develop into an adult. Fully-grown caterpillars tend to be green. After hatching, the caterpillar usually eats the egg shell. The caterpillars have been known to eat from the
black cherry ''Prunus serotina'', commonly called black cherry,World Economic Plants: A Standard Reference, Second Edition'. CRC Press; 19 April 2016. . p. 833–. wild black cherry, rum cherry, or mountain black cherry, is a deciduous tree or shrub of the g ...
.


Ecology


Life cycle

The Appalachian tiger swallowtail is univoltine. Female butterflies usually lay their singular brood from May to June. During the mating season, the males fly through the treetops, scouting for females. Females, though, tend to be difficult to find because they prefer to hide in the forest canopy. After mating, females lay their eggs singularly on host plant leaves. The caterpillars, after hatching, then eat the host plants' leaves and undergo
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 ...
throughout the winter.
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 ...
tion occurs around late June to July and pupae
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
until spring. Adults hatch in May.


Habitat

Appalachian tiger swallowtail butterflies tend to congregate in the woodlands and woodland openings. They prefer to remain in mid-level to high elevations in the southern Appalachian Mountains, and the species' range stretches from Pennsylvania to Georgia and Alabama. This species is extremely prevalent at higher elevations, but can also be found at lower elevations or valleys within mountainous terrain. The adult butterflies tend to remain in the canopy of the Appalachian hardwood forest and mainly only leave when visiting flowers. Their habitats often parallel those of ''P. glaucus''.


Nectar plants

Adults drink nectar from many plants, including the mountain laurel (''
Kalmia latifolia ''Kalmia latifolia'', the mountain laurel, calico-bush, or spoonwood, is a species of flowering plant in the heath family Ericaceae, that is native to the eastern United States. Its range stretches from southern Maine south to northern Florida, ...
''), the pink azalea (''
Rhododendron ''Rhododendron'' (; from Ancient Greek ''rhódon'' "rose" and ''déndron'' "tree") is a very large genus of about 1,024 species of woody plants in the heath family (Ericaceae). They can be either evergreen or deciduous. Most species are nati ...
nudiflorum''), the blackberry (''
Rubus ''Rubus'' is a large and diverse genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae, with over 1,350 species. Raspberries, blackberries, and dewberries are common, widely distributed members of the genus. Most of these ...
''), the cockspur thorn (''
Crataegus crus-galli ''Crataegus crus-galli'' is a species of hawthorn known by the common names cockspur hawthorn and cockspur thorn. It is native to eastern North America from Ontario to Texas to Florida, and it is widely used in horticulture. It is thought to be th ...
''), the multiflora rose (''
Rosa multiflora ''Rosa multiflora'' — ( syn. ''Rosa polyantha'') is a species of rose known commonly as multiflora rose, baby rose, Japanese rose, many-flowered rose, seven-sisters rose, Eijitsu rose and rambler rose. It is native to eastern Asia, in China, ...
''), and the bush honeysuckle (''
Diervilla ''Diervilla'', or bush honeysuckle, is a genus of three species of deciduous shrubs in the family Caprifoliaceae, all indigenous to eastern North America. The genus is named after a French surgeon Dr. Marin Diereville, who introduced the plant ...
''). Near the summit, they also tend to drink from the autumn olive ('' Elaeagnus commutata''), but at lower elevation, they drink from the daisy fleabane (''Erigeron ramosus'').


Host plants

The larvae eat the leaves of a wide variety of trees, including cottonwood, tulip tree, lemon and cherry.


Flight patterns

The Appalachian tiger swallowtail shares a delayed flight pattern and many other hybrid-like traits with ''P. glaucus'' and ''P. canadensis'' despite its genetic isolation. Its flight period is in May. Some individuals take flight during late June, while others fly from May to early June. Populations of this species make annual migrations of over . ''Papilio appalachiensis'' usually has a singular flight period from spring to early summer and emerges later than ''P. glaucus''. The Appalachian tiger swallowtail tends to fly directly up the western side of
Spruce Knob Spruce Mountain, located in eastern West Virginia, is the highest ridge of the Allegheny Mountains. The whale-backed ridge extends for only from northeast to southwest, but several of its peaks exceed in elevation. The summit, Spruce Knob (4863 ...
towards the summit against strong westerly winds. The summit contains substantial vegetation within protected nooks where the adult butterflies rest and obtain nectar from various plants.


References

{{Taxonbar, from=Q744621 appalachiensis Butterflies of North America Hybrid animals Butterflies described in 2002