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{{Short description|Species of butterfly}}
{{Short description|Species of butterfly}}
{{Speciesbox
{{italic title}}
{{Taxobox
| name = Silver-spotted skipper
| name = Silver-spotted skipper
| image = Silver-spotted Skipper Epargyreus clarus Feeding 2116px.jpg
| image = Silver-spotted Skipper Epargyreus clarus Feeding 2116px.jpg
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| ordo = [[Lepidoptera]]
| familia = [[Hesperiidae]]
| genus = ''[[Epargyreus]]''
| species = '''''E. clarus'''''
| binomial = ''Epargyreus clarus''
| binomial_authority = ([[Pieter Cramer|Cramer]], 1775)
| range_map = E. clarus map.jpeg
| status = G5
| status = G5
| status_system = TNC
| status_system = TNC
| status_ref = <ref>{{cite web |title=NatureServe Explorer 2.0 Epargyreus clarus Silver-spotted Skipper |url=https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.118198/Epargyreus_clarus |website=explorer.natureserve.org |access-date=1 October 2020}}</ref>
| status_ref = <ref>{{cite web |title=NatureServe Explorer 2.0 Epargyreus clarus Silver-spotted Skipper |url=https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.118198/Epargyreus_clarus |website=explorer.natureserve.org |access-date=1 October 2020}}</ref>
| taxon = Epargyreus clarus
| tribus = [[Eudamini]]
| authority = ([[Pieter Cramer|Cramer]], 1775)
| synonyms =
| range_map = E. clarus map.jpeg
| synonyms = {{Specieslist
|Papilio tityrus|Fabricius, 1775
|Papilio clarus|Cramer, [1775]
|Epargyreus tityrus|(Fabricius, 1775)
|Epargyreus titgrus|Draudt, 1922 (Mis-spelling)
}}
| synonyms_ref =
}}
}}


'''''Epargyreus clarus''''', the '''silver-spotted skipper''', is a [[butterfly]] of the family [[Hesperiidae]]. It is claimed to be the most recognized skipper in [[North America]].<ref>{{cite book |last=Kaufman |first= Kenn |author-link= Kenn Kaufman |author2=Eaton, Eric R. |title= Kaufman Field Guide to Insects of North America |year= 2007 |publisher= Houghton Mifflin Books |isbn= 978-0-618-15310-7 |pages= 236–237}}</ref> ''E. clarus'' occurs in fields, gardens, and at forest edges and ranges from southern [[Canada]] throughout most of the [[United States]] to northern [[Mexico]], but is absent in the [[Great Basin]] and western [[Texas]].<ref name="BAMONA">{{cite web|url=http://www.butterfliesandmoths.org/species?l=1905|title=Silver-spotted skipper|publisher=Butterflies and Moths of North America|access-date=2008-10-11|archive-url=https://web.archive.org/web/20090116111214/http://www.butterfliesandmoths.org/species?l=1905|archive-date=2009-01-16|url-status=dead}}</ref>
'''''Epargyreus clarus''''', the '''silver-spotted skipper''', is a [[butterfly]] of the family [[Hesperiidae]]. It is claimed to be the most recognized skipper in North America.<ref>{{cite book |last=Kaufman |first= Kenn |author-link= Kenn Kaufman |author2=Eaton, Eric R. |title= Kaufman Field Guide to Insects of North America |year= 2007 |publisher= Houghton Mifflin Books |isbn= 978-0-618-15310-7 |pages= 236–237}}</ref> {{Nowrap|''E. clarus''}} occurs in fields, gardens, and at forest edges and ranges from southern Canada throughout most of the United States to northern Mexico, but is absent in the [[Great Basin]] and western [[Texas]].<ref name="BAMONA">{{cite web|url=http://www.butterfliesandmoths.org/species?l=1905|title=Silver-spotted skipper|publisher=Butterflies and Moths of North America|access-date=2008-10-11|archive-url=https://web.archive.org/web/20090116111214/http://www.butterfliesandmoths.org/species?l=1905|archive-date=2009-01-16|url-status=dead}}</ref>


''E. clarus'' larvae create and reside in unique shelters stuck together with [[silk]], which do not protect them from predators.<ref name=":3" /> Natural predators of the species include paper wasp foragers, sphecid wasp and ''Crematogaster opuntiae'' ants.<ref name=":2" /><ref name=":4" /> The species is also considered to be a perching species, meaning that adult males compete for territory to attract females.<ref name=":5">{{Cite journal|last=Scott|first=James A.|date=1974|title=Mate-Locating Behavior of Butterflies|journal=The American Midland Naturalist|volume=91|issue=1|pages=103–117|doi=10.2307/2424514|jstor=2424514}}</ref> Although ''E. clarus'' is considered to be a pest of a few crop plants such as beans, its pest activity is not serious enough to warrant initiating major control measures.<ref name=":2" />
{{Nowrap|''E. clarus''}} larvae create and reside in unique shelters stuck together with [[silk]], which do not protect them from predators.<ref name=":3" /> Natural predators of the species include paper wasp foragers, sphecid wasp and ''Crematogaster opuntiae'' ants.<ref name=":2" /><ref name=":4" /> The species is also considered to be a perching species, meaning that adult males compete for territory to attract females.<ref name=":5">{{Cite journal|last=Scott|first=James A.|date=1974|title=Mate-Locating Behavior of Butterflies|journal=The American Midland Naturalist|volume=91|issue=1|pages=103–117|doi=10.2307/2424514|jstor=2424514}}</ref> Although {{Nowrap|''E. clarus''}} is considered to be a pest of a few crop plants such as beans, its pest activity is not serious enough to warrant initiating major control measures.<ref name=":2" />
[[File:Silver-spotted Skipper - Hodges-3870 (Epargyreus clarus).jpg|alt=Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Caterpillar|thumb|Silver-spotted skipper - Hodges#3870 (''Epargyreus clarus'') Caterpillar feeding on Wisteria]]
[[File:Silver-spotted Skipper (Epargyreus clarus) .jpg|alt=Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Feeding on Butterfly Bush|thumb|Silver-spotted skipper - Hodges#3870 (''Epargyreus clarus'') Feeding on Butterfly Bush]]


== Geographic range ==
== Geographic range ==
''E. clarus'' has a wide distribution throughout [[North America]]: it ranges from southern Canada throughout most of the United States to northern Mexico, but is absent from the [[Great Basin]] and west Texas.<ref name="BAMONA" />
''Epargyreus clarus'' has a wide distribution throughout North America: it ranges from southern Canada throughout most of the United States to northern Mexico, but is absent from the Great Basin and western Texas.<ref name="BAMONA" />


== Habitat ==
== Habitat ==
[[File:Epargyreus clarus on wooden floor.jpeg|thumb|Sliver-spotted skipper on a wooden floor at Morgan Falls Park in [[Sandy Springs, Georgia]].]]
The silver-spotted skipper prefers open ranges where nectar plants are found, such as forest edges, swamps, brushy areas, and [[Riparian zone|riparian]] habitats at lower elevations.<ref name=":0">{{Cite journal|last=Shapiro|first=Arthur M.|date=1978|title=Phenotypic and Behavioral Convergence of "Silver-Spotted Skippers" (Lepidoptera: Hesperiidae)|journal=Biotropica|volume=10|issue=2|pages=159–160|doi=10.2307/2388021|jstor=2388021}}</ref><ref name=":1">{{Cite book|title=LEPIDOPTERA OF THE PACIFIC NORTHWEST: CATERPILLARS AND ADULTS|last1=Miller|first1=Jeffrey C.|last2=Hammond|first2=Paul C.|year=2003|pages=28}}</ref>
The silver-spotted skipper prefers open ranges where nectar plants are found, such as forest edges, swamps, brushy areas, and [[Riparian zone|riparian]] habitats at lower elevations.<ref name=":0">{{Cite journal|last=Shapiro|first=Arthur M.|date=1978|title=Phenotypic and Behavioral Convergence of "Silver-Spotted Skippers" (Lepidoptera: Hesperiidae) |journal=Biotropica |volume=10 |issue=2 |pages=159–160 |doi=10.2307/2388021 |jstor=2388021|bibcode=1978Biotr..10..159S }}</ref><ref name=":1">{{Cite book |title=LEPIDOPTERA OF THE PACIFIC NORTHWEST: CATERPILLARS AND ADULTS |last1=Miller|first1=Jeffrey C.|last2=Hammond|first2=Paul C.|year=2003|pages=28}}</ref>


== Home range ==
== Home range ==
At night, or on hot or cloudy days, ''E. clarus'' rests on the underside of a leaf. It hangs upside down, holding its wings together over its back so that its silver spot is exposed. ''E. clarus'' is often restricted by its habitat, preferring to fly in the shade.<ref name=":0" />
At night, or on hot or cloudy days, ''Epargyreus clarus'' rests on the underside of a leaf. It hangs upside down, holding its wings together over its back so that its silver spot is exposed. {{Nowrap|''E. clarus''}} is often restricted by its habitat, preferring to fly in the shade.<ref name=":0" />


== Food resources ==
== Food resources ==


=== Caterpillars ===
=== Caterpillars ===
''E. clarus'' caterpillars consume leaves of herbs, vines, shrubs, and trees in the pea family (''[[Fabaceae]]'') including false indigobush (''[[Amorpha fruticosa]]''), American hogpeanut (''[[Amphicarpaea bracteata]]''), groundnut (''[[Apios americana]]''), American wisteria (''[[Wisteria frutescens]]''), Atlantic pigeonwings or butterfly pea (''[[Clitoria mariana]]''), and the introduced Dixie ticktrefoil (''Desmodium tortuosum''), kudzu (''[[Pueraria montana]]''), black locust (''[[Robinia pseudoacacia]]''), Chinese wisteria (''[[Wisteria sinensis]]'') and an assortment of other legumes.<ref name=":1" />
''Epargyreus clarus'' caterpillars consume leaves of herbs, vines, shrubs, and trees in the pea family (''[[Fabaceae]]'') including false indigobush (''[[Amorpha fruticosa]]''), American hogpeanut (''[[Amphicarpaea bracteata]]''), groundnut (''[[Apios americana]]''), American wisteria (''[[Wisteria frutescens]]''), Atlantic pigeonwings or butterfly pea (''[[Clitoria mariana]]''), and the introduced Dixie ticktrefoil (''[[Desmodium tortuosum]]''), kudzu (''[[Pueraria montana]]''), black locust (''[[Robinia pseudoacacia]]''), Chinese wisteria (''[[Wisteria sinensis]]'') and an assortment of other legumes.<ref name=":1" />


Innate host plant preferences confer greater performance on larvae, due to differences in leaf nutrient concentrations. However, preference may be influenced by a larva’s rearing host species.<ref>{{Cite journal|last1=Rosenwald|first1=Laura C.|last2=Lill|first2=John T.|last3=Lind|first3=Eric M.|last4=Weiss|first4=Martha R.|date=2017-05-10|title=Dynamics of host plant selection and host-switching by silver-spotted skipper caterpillars|journal=Arthropod-Plant Interactions|language=en|pages=833–842|doi=10.1007/s11829-017-9538-0|issn=1872-8855|volume=11|issue=6|s2cid=37474805}}</ref>
Innate host plant preferences confer greater performance on larvae, due to differences in leaf nutrient concentrations. However, preference may be influenced by a larva’s rearing host species.<ref>{{Cite journal|last1=Rosenwald|first1=Laura C.|last2=Lill|first2=John T.|last3=Lind|first3=Eric M.|last4=Weiss|first4=Martha R.|date=2017-05-10|title=Dynamics of host plant selection and host-switching by silver-spotted skipper caterpillars|journal=Arthropod-Plant Interactions|language=en|pages=833–842|doi=10.1007/s11829-017-9538-0|issn=1872-8855|volume=11|issue=6|bibcode=2017APInt..11..833R |s2cid=37474805}}</ref>

They live in meadows and swamps. Their average size is 2 inches {{Nowrap|(5 cm)}} long. Average life span is about 2 months. Their predators are wasps, ants, birds, frogs and spiders.<ref>Melissa Stewart, National Geographic Kids, Ick!, Washington DC, Text Copyright 2020</ref>


=== Adults ===
=== Adults ===
Adults use their long tongues to feed on the nectar of a variety of flowers, mud, and even animal feces.<ref name=":2">{{Cite journal|last=Hall|first=Donald W.|date=2017|title=Silver-Spotted Skipper Epargyreus clarus (Cramer)|journal=The Institute of Food and Agricultural Sciences Extension}}</ref> They almost never visit yellow flowers, favoring blue, red, pink, purple, and sometimes white and cream-colored ones. These include everlasting pea, common milkweed, red clover, buttonbush, blazing star, and thistles.<ref name="BAMONA" />
Adults use their long tongues to feed on the nectar of a variety of flowers, mud, and animal feces.<ref name=":2">{{Cite journal|last=Hall|first=Donald W.|date=2017|title=Silver-Spotted Skipper Epargyreus clarus (Cramer)|journal=The Institute of Food and Agricultural Sciences Extension}}</ref> They almost never visit yellow flowers, favoring those that are blue, red, pink, purple, and sometimes white and cream. These include everlasting pea, common milkweed, red clover, buttonbush, blazing star, and thistles.<ref name="BAMONA" />


== Parental care ==
== Parental care ==
=== Oviposition ===
=== Oviposition ===
Females lay a single egg on or occasionally nearby host plants. They have been reported to oviposit on the least snoutbean (''[[Rynchosia minima]]'').<ref>{{Cite book|title=The Butterflies of North America: A Natural History and Field Guide.|last=Scott|first=JA|publisher=Stanford University Press|year=1986|pages=583}}</ref>
Females lay a single egg on or occasionally near host plants. They have been reported to oviposit on the least snoutbean (''[[Rynchosia minima]]'').<ref>{{Cite book|title=The Butterflies of North America: A Natural History and Field Guide.|last=Scott|first=JA|publisher=Stanford University Press|year=1986|pages=583}}</ref>


== Life cycle ==
== Life cycle ==
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====Shelter building====
====Shelter building====
All larval instars of ''E. clarus'' build shelters on their host plant. They build about 5 shelters throughout larval growth and development in 4 distinct styles, which are constructed from instinct and unique to larval size. The first, second, and third instars make a simpler, invariant structure that requires 2 incisions in the leaf and silk to fold over the flap created. Shelters are typically built on the apical portion of the leaflet. One study of ''E. clarus'' larvae in the Washington D.C. area defined 5 steps of shelter building:<ref name=":3">{{Cite journal|last1=Weiss|first1=Martha R.|last2=Lind|first2=Eric M.|last3=Jones|first3=Meg T.|last4=Long|first4=Jeremy D.|last5=Maupin|first5=Jennifer L.|date=2003-07-01|title=Uniformity of Leaf Shelter Construction by Larvae of Epargyreus clarus (Hesperiidae), the Silver-Spotted Skipper|journal=Journal of Insect Behavior|language=en|volume=16|issue=4|pages=465–480|doi=10.1023/A:1027399122333|s2cid=28669353|issn=0892-7553}}</ref>
All larval instars of ''Epargyreus clarus'' build shelters on their host plant. They build about five shelters throughout larval growth and development in four distinct styles. These are constructed from instinct and are unique to larval size. The first, second, and third instars make a simple, invariant structure that requires two incisions in the leaf and silk to fold over the flap created. Shelters are typically built on the apical portion of the leaflet. One study of {{Nowrap|''E. clarus''}} larvae in the Washington D.C. area defined five steps of shelter building:<ref name=":3">{{Cite journal|last1=Weiss|first1=Martha R.|last2=Lind|first2=Eric M.|last3=Jones|first3=Meg T.|last4=Long|first4=Jeremy D.|last5=Maupin|first5=Jennifer L.|date=2003-07-01|title=Uniformity of Leaf Shelter Construction by Larvae of Epargyreus clarus (Hesperiidae), the Silver-Spotted Skipper|journal=Journal of Insect Behavior|language=en|volume=16|issue=4|pages=465–480|doi=10.1023/A:1027399122333|s2cid=28669353|issn=0892-7553}}</ref>
# ''Site selection'': Larvae traverse across the entire leaf, following major veins and swinging their heads to create a single silk trail.
# ''Site selection'': Larvae traverse across the entire leaf, following major veins and swinging their heads to create a single silk trail.
# Silk template: A thick, visible mat of silk outlines the intended incisions and ultimately becomes the ceiling of the leaf house. The larvae keep their posterior end on the template at all times.
# Silk template: A thick, visible mat of silk outlines the intended incisions and ultimately becomes the ceiling of the leaf house. The larvae keep their posterior end on the template at all times.
# ''First cut'': The first cut follows the silk template on the perimeter nearest the leaf apex. The larvae often interrupt chewing cuts in order to deposit silk at the terminus. The first cut is completed in about 6-7 segments. At the end, the larva makes a U-turn motion so that its position relative to the leaf surface and template margins remains the same while facing the opposite direction.
# ''First cut'': The first cut follows the silk template on the perimeter nearest the leaf apex. The larvae often interrupt chewing cuts in order to deposit silk at the terminus. The first cut is completed in about 6 to 7 segments. At the end, the larva makes a U-turn motion so that its position relative to the leaf surface and template margins remains the same while facing the opposite direction.
# ''Second cut and notch'': The second cut follows the silk template on the edge closest to the base of the leaflet and is completed in about 2 to 4 segments. The first segment curves inwards, is made in a single effort, and is about 75% the length of the first cut. The next segment is adjacent to the initial segment and continues past the length of the first cut. The juncture between the two segments creates a notch.
# ''Second cut and notch'': The second cut follows the silk template on the edge closest to the base of the leaflet and is completed in about 2 to 4 segments. The first segment curves inwards, is made in a single effort and is about 75% the length of the first cut. The next segment is adjacent to the initial segment and continues past the length of the first cut. The juncture between the two segments creates a notch.
# ''Folding and securing'': The uncut area between the first and second incisions composes the hinge. The hinge length tends to correspond to larval body length. The larva lay silk strands along this area, parallel to the cut termini. As the strands contract, the notch creates a pinch and the leaf flap is pulled over the leaflet plane. The pinch then becomes the peak of the roof and the larva begins to make “guy-wires” that attach the edge of the flap to the leaflet surface.
# ''Folding and securing'': The uncut area between the first and second incisions composes the hinge. The hinge length tends to correspond to larval body length. The larvae lay silk strands along this area, parallel to the cut termini. As the strands contract, the notch creates a pinch, and the leaf flap is pulled over the leaflet plane. The pinch then becomes the peak of the roof and the larva begins to make "guy-wires" that attach the edge of the flap to the leaflet surface.
Larvae take breaks of about 30 minutes between each cut and fold. It typically takes less than 2 and a half hours to complete shelter construction. Larvae spend about 95% of their time resting on the ceiling of the shelter.<ref name=":3" />
Larvae take breaks of about 30 minutes between each cut and fold. It typically takes less than 2{{half}} hours to complete shelter construction. Larvae spend about 95% of their time resting on the ceiling of the shelter.<ref name=":3" />
[[File:Pinned E. clarus.jpg|thumb|right|Adult]]
[[File:Pinned E. clarus.jpg|thumb|right|Adult]]
[[File:Silver spotted skipper.webm|thumb|thumbtime=7|right|[[Epargyreus clarus|the silver-spotted skipper]] [[nectar|nectaring]]]]
First instar shelters are typically fastened by only about 2-4 “guy-wires. In addition, older caterpillars occasionally live in a nest made of multiple leaves connected by silk, especially when using host plants with smaller leaflets. For example, first, second, and third instars can make shelters with one ''Robinia'' leaflet, but the fourth and fifth instars have to use 2 to 4 or 5 leaflets.<ref name=":3" />

First instar shelters are typically fastened by only about 2 to 4 "guy-wires". In addition, older caterpillars occasionally live in a nest made of multiple leaves connected by silk, especially when using host plants with smaller leaflets. For example, first, second, and third instars can make shelters with one ''Robinia'' leaflet, but the fourth and fifth instars have to use 2–4 or 5 leaflets.<ref name=":3" />


=== Pupa ===
=== Pupa ===
Larvae pupate throughout the winter. The pupa is dark brown with black and white marks. There are three or more overlapping generations of hibernating pupa in Florida – fewer in the northern regions.<ref name=":2" />
Larvae of ''Epargyreus clarus'' pupate throughout the winter. The pupa is dark brown with black and white marks. There are three or more overlapping generations of hibernating pupa in Florida – fewer in the northern regions.<ref name=":2" />


=== Adult ===
=== Adult ===
The adult wingspan ranges from 1.75 to 2.625 inches (4.5 to 6.7&nbsp;cm). Each forewing is triangular and dark brown with a large yellow-orange medial patch. Male forewings are more pointed than those of females. The hindwings are also dark brown, with a large silver patch on the distal third of the ventral side.<ref name=":1" /><ref name=":2" />
The adult wingspan ranges from 1.75 to 2.625 inches (4.5 to 6.7&nbsp;cm). Each forewing is triangular and dark brown with a large yellow-orange medial patch. Male forewings are more pointed than those of females. The hindwings are also dark brown, with a large silver patch on the discal third of the ventral side.<ref name=":1" /><ref name=":2" />


== Migration ==
== Migration ==
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=== Predators ===
=== Predators ===
[[File:Wheel bug assassin bug vs. silver-spotted skipper caterpillar.webm|Wheel bug assassin bug vs. silver-spotted skipper caterpillar|thumb|right]]
[[File:Wheel bug assassin bug vs. silver-spotted skipper caterpillar.webm|Wheel bug assassin bug vs. silver-spotted skipper caterpillar|thumb|right]]Common predators include the paper wasp foragers ''[[Polistes fuscatus]]'' and ''[[Polistes dominula]]'', as well as ''Crematogaster opuntiae'' ants. Studies have found that shelters protect larvae from these predators in lab settings. Shelters delayed or prevented ''Crematogaster opuntiae'' colonies from detecting the larvae. This is likely because of these ants’ limited senses, in conjunction with the inhibitory structure of the shelter: they are able to detect a motionless caterpillar only within approximately one body length, and this is further obstructed by the single small opening into the shelter. While wasps in the laboratory setting did spend more time on areas of the leaf damaged by feeding or silk deposition, the larval shelter prevented visual detection and posed a physical barrier. However, shelters did not protect against these predators in the field. Wasps need spend only a few seconds on a leaflet to successfully identify and remove the larva from its shelter. This suggests that shelter identification and larval extraction is a learned ability, perhaps developed through visual or olfactory cues.<ref name=":4">{{Cite journal|last1=Jones|first1=Meg T.|last2=Castellanos|first2=Ignacio|last3=Weiss|first3=Martha R.|date=2002-12-01|title=Do leaf shelters always protect caterpillars from invertebrate predators?|journal=Ecological Entomology|language=en|volume=27|issue=6|pages=753–757|doi=10.1046/j.1365-2311.2002.00465.x|s2cid=86245610|issn=1365-2311}}</ref>
Common predators of ''Epargyreus clarus'' include the paper wasp foragers ''[[Polistes fuscatus]]'' and ''[[Polistes dominula]]'', as well as ''Crematogaster opuntiae'' ants. Studies have found that shelters protect larvae from these predators in lab settings. Shelters delayed or prevented ''Crematogaster opuntiae'' colonies from detecting the larvae. This is likely because of the ants' limited senses, in conjunction with the inhibitory structure of the shelter: they are able to detect a motionless caterpillar only within approximately one body length, and this is further obstructed by the single small opening into the shelter. While wasps in the laboratory setting did spend more time on areas of the leaf damaged by feeding or silk deposition, the larval shelter prevented visual detection and posed a physical barrier. However, shelters did not protect against these predators in the field. Wasps need spend only a few seconds on a leaflet to successfully identify and remove the larva from its shelter. This suggests that shelter identification and larval extraction is a learned ability, perhaps developed through visual or olfactory cues.<ref name=":4">{{Cite journal|last1=Jones|first1=Meg T.|last2=Castellanos|first2=Ignacio|last3=Weiss|first3=Martha R.|date=2002-12-01|title=Do leaf shelters always protect caterpillars from invertebrate predators? |journal=Ecological Entomology |language=en |volume=27 |issue=6 |pages=753–757 |doi=10.1046/j.1365-2311.2002.00465.x |bibcode=2002EcoEn..27..753J |s2cid=86245610 |issn=1365-2311}}</ref>


Other predators include the sphecid wasp ''[[Horse guard wasp|Stictia carolina]]'', which sometimes supplies its nests with silver-spotted skipper larvae. Other predators also often are attracted to the chemical cues from [[frass]], or insect excrement. ''E. clarus'' larvae use an anal comb to throw their frass up to 38 body lengths away from them. When disturbed, larvae regurgitate an unpleasant, greenish chemical in defense.<ref name=":2" />
Other predators include the sphecid wasp ''[[Horse guard wasp|Stictia carolina]]'', which sometimes supplies its nests with silver-spotted skipper larvae. Additional predators also often are attracted to the chemical cues from [[frass]], or insect excrement. ''Epargyreus clarus'' larvae use an anal comb to throw their frass up to 38 body lengths away from them. When disturbed, larvae regurgitate an unpleasant, greenish chemical in defense.<ref name=":2" />


=== Parasites ===
=== Parasites ===
''E. clarus'' larvae are also a natural host of ''[[Baculoviridae|Baculovirus]]''. The advanced stages of this infection cause larvae to feed without resting. Later, they climb to higher and more exposed areas, where they are more conspicuous to predators. Predator attack leaves their infected remains to be absorbed by the surrounding vegetation, which will be consumed by new hosts.<ref name=":2" />
{{Nowrap|''E. clarus''}} larvae are a natural host of ''[[Baculoviridae|Baculovirus]]''. The advanced stages of this infection cause larvae to feed without resting. Later, they climb to higher and more exposed areas, where they are more conspicuous to predators. Predator attack leaves their infected remains to be absorbed by the surrounding vegetation, which will be consumed by new hosts.<ref name=":2" />


== Mating ==
== Mating ==
Instead of receiving ejaculate, a female receives a large [[spermatophore]] from a male for each copulation. Thus, the number of spermatophores a female contains roughly equates to the number of copulations. Generally, when there are more than two spermatophores present in a female, each one’s placement and degree of erosion indicates its age relative to the other. ''E. clarus'', however, is particularly resistant to erosion. Once mated, females seem to reject males, at least for some time afterwards.<ref>{{Cite journal|last=Burns|first=John M.|date=1968|title=Mating Frequency in Natural Populations of Skippers and Butterflies as Determined by Spermatophore Counts|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=61|issue=3|pages=852–859|doi=10.1073/pnas.61.3.852|pmid=16591715|jstor=58962|pmc=305406}}</ref>
Instead of receiving ejaculate, a female receives a large [[spermatophore]] from a male for each copulation. Thus, the number of spermatophores a female contains roughly equates to the number of copulations. Generally, when there are more than two spermatophores present in a female, each one’s placement and degree of erosion indicates its age relative to the other. ''Epargyreus clarus'' is particularly resistant to erosion. Once mated, females seem to reject males, at least for some time afterwards.<ref>{{Cite journal|last=Burns|first=John M.|date=1968|title=Mating Frequency in Natural Populations of Skippers and Butterflies as Determined by Spermatophore Counts |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=61 |issue=3 |pages=852–859 |doi=10.1073/pnas.61.3.852 |pmid=16591715 |jstor=58962 |pmc=305406|bibcode=1968PNAS...61..852B|doi-access=free}}</ref>


=== Male defense of places likely to attract females ===
=== Male defense of places likely to attract females ===
Males perch on branches and tall weeds about 1–2 m above the ground, darting out when any insect passes in hopes of finding a receptive ''E. clarus'' female. When confronted by another conspecific, male movements are swift and acrobatic. This perching behavior is part of a common mate-location strategy, in which males compete for places where females are more likely to occur.<ref name=":5" /> Females do not engage in this behavior, but may have to fly considerable distances to find mates, depending on the proximity of male territory to their larval development site. The places "defended" by males are roughly the size of a small room and are most likely chosen based on vegetation and topography.<ref name=":0" />
Males perch on branches and tall weeds about {{Convert|1|to|2|m|ft|order=flip}} above the ground, darting out when any insect passes in hopes of finding a receptive ''Epargyreus clarus'' female. When confronted by another conspecific, male movements are swift and acrobatic. This perching behavior is part of a common mate-location strategy in which males compete for places where females are more likely to occur.<ref name=":5" /> Females do not engage in this behavior, but may have to fly considerable distances to find mates, depending on the proximity of male territory to their larval development site. The places "defended" by males are roughly the size of a small room and are most likely chosen based on vegetation and topography.<ref name=":0" />


== Physiology ==
== Physiology ==

=== Vision ===
=== Vision ===
''E. clarus'' possesses [[compound eyes]] that lack pigment in the iris region. Each [[Ommatidium|ommatidia]], or single optical unit, has its own unique visual field that spans about 2°. The small visual field can be due to crystalline tracts in the eye that restrict the light to reach the retina only through this path.<ref>{{cite journal | last1 = Doving | first1 = K. B. | year = 1969| title = Function of Insect Compound Eyes Containing Crystalline Tracts | journal = The Journal of General Physiology | volume = 54 | issue = 2| pages = 250–267 | doi = 10.1085/jgp.54.2.250 | pmid = 4894303 | pmc = 2225925 | citeseerx = 10.1.1.282.9479 }}</ref>
''Epargyreus clarus'' possesses [[compound eyes]] that lack pigment in the iris region. Each [[Ommatidium|ommatidia]], or single optical unit, has its own unique visual field that spans about 2°. The small visual field can be due to crystalline tracts in the eye that restrict the light to reach the retina only through this path.<ref>{{cite journal | last1 = Doving | first1 = K. B. | year = 1969| title = Function of Insect Compound Eyes Containing Crystalline Tracts | journal = The Journal of General Physiology | volume = 54 | issue = 2| pages = 250–267 | doi = 10.1085/jgp.54.2.250 | pmid = 4894303 | pmc = 2225925 | citeseerx = 10.1.1.282.9479 }}</ref>


== Interaction with plants ==
== Interaction with plants ==
[[File:Silver Spotted Skpper (6231641469).jpg|thumb|Silver Spotted Skipper nectaring ''Liatris'']]
[[File:Silver Spotted Skpper (6231641469).jpg|thumb|Silver-spotted skipper nectaring ''Liatris'']]
''E. clarus'' is known to be a nectar-thief species. An individual nectar thief is an animal that takes nectar from a flower but does not pollinate it while doing so. A species as a whole can be considered a thief species if it does not pollinate a species of flowers during more than 50% of its feedings. Research has found that while foraging on flowers, the butterfly tended to probe the innermost disk florets, which are the male organs, but not make contact with the outermost florets, which are the female organs.<ref>{{Cite journal|last1=Venables|first1=Adrienne|last2=Barrows|first2=Edward|date=1985|title=SKIPPERS: POLLINATORS OR NECTAR THIEVES?|url=http://images.peabody.yale.edu/lepsoc/jls/1980s/1985/1985-39(4)299-Venables.pdf|journal=Journal of the Lepidopterists' Society|volume=39 | issue = 4 |pages=299–312}}</ref>
''Epargyreus clarus'' is known to be a nectar-thief species. An individual nectar thief is an animal that takes nectar from a flower but does not pollinate it while doing so. A species as a whole can be considered a thief species if it does not pollinate a species of flowers during more than 50% of its feedings. Research has found that while foraging on flowers, the butterfly tended to probe the innermost disk florets, which are the male organs, but not make contact with the outermost florets, which are the female organs.<ref>{{Cite journal|last1=Venables|first1=Adrienne|last2=Barrows|first2=Edward|date=1985 |title=SKIPPERS: POLLINATORS OR NECTAR THIEVES? |url=http://images.peabody.yale.edu/lepsoc/jls/1980s/1985/1985-39(4)299-Venables.pdf |journal=Journal of the Lepidopterists' Society |volume=39 | issue = 4 |pages=299–312}}</ref>


== Pest of crop plants ==
== Pest of crop plants ==
''E. clarus'' is known to feed on various crop plants such as soybean and kidney bean. However, no large-scale control measures have been taken as the species' pest activity is not too detrimental.<ref name=":2" />
''Epargyreus clarus'' is known to feed on various crop plants such as soybean and kidney bean. However, no large-scale control measures have been taken, as the species' pest activity is not too detrimental.<ref name=":2" />


==References==
==References==
Line 114: Line 118:
{{Commonscat}}
{{Commonscat}}
* [http://entomology.ifas.ufl.edu/creatures/bfly/silver-spotted_skipper.htm ''Epargyreus clarus'', silver-spotted skipper] on the [[University of Florida|UF]] / [[Institute of Food and Agricultural Sciences|IFAS]] Featured Creatures Web site
* [http://entomology.ifas.ufl.edu/creatures/bfly/silver-spotted_skipper.htm ''Epargyreus clarus'', silver-spotted skipper] on the [[University of Florida|UF]] / [[Institute of Food and Agricultural Sciences|IFAS]] Featured Creatures Web site

{{Taxonbar|from=Q568157}}
{{Taxonbar|from=Q568157}}


[[Category:Eudaminae]]
[[Category:Butterflies of North America]]
[[Category:Butterflies of North America]]
[[Category:Articles containing video clips]]
[[Category:Articles containing video clips]]
[[Category:Eudaminae]]
[[Category:Butterflies described in 1775]]
[[Category:Butterflies described in 1775]]
[[Category:Taxa named by Pieter Cramer]]
[[Category:Taxa named by Pieter Cramer]]
[[Category:Lepidoptera of the United States]]
[[Category:Lepidoptera of Mexico]]
[[Category:Lepidoptera of Canada]]

Latest revision as of 06:43, 25 September 2024

Silver-spotted skipper

Secure  (NatureServe)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Hesperiidae
Genus: Epargyreus
Species:
E. clarus
Binomial name
Epargyreus clarus
(Cramer, 1775)
Synonyms
  • Papilio tityrus Fabricius, 1775
  • Papilio clarus Cramer, [1775]
  • Epargyreus tityrus (Fabricius, 1775)
  • Epargyreus titgrus Draudt, 1922 (Mis-spelling)

Epargyreus clarus, the silver-spotted skipper, is a butterfly of the family Hesperiidae. It is claimed to be the most recognized skipper in North America.[2] E. clarus occurs in fields, gardens, and at forest edges and ranges from southern Canada throughout most of the United States to northern Mexico, but is absent in the Great Basin and western Texas.[3]

E. clarus larvae create and reside in unique shelters stuck together with silk, which do not protect them from predators.[4] Natural predators of the species include paper wasp foragers, sphecid wasp and Crematogaster opuntiae ants.[5][6] The species is also considered to be a perching species, meaning that adult males compete for territory to attract females.[7] Although E. clarus is considered to be a pest of a few crop plants such as beans, its pest activity is not serious enough to warrant initiating major control measures.[5]

Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Caterpillar
Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Caterpillar feeding on Wisteria
Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Feeding on Butterfly Bush
Silver-spotted skipper - Hodges#3870 (Epargyreus clarus) Feeding on Butterfly Bush

Geographic range

[edit]

Epargyreus clarus has a wide distribution throughout North America: it ranges from southern Canada throughout most of the United States to northern Mexico, but is absent from the Great Basin and western Texas.[3]

Habitat

[edit]
Sliver-spotted skipper on a wooden floor at Morgan Falls Park in Sandy Springs, Georgia.

The silver-spotted skipper prefers open ranges where nectar plants are found, such as forest edges, swamps, brushy areas, and riparian habitats at lower elevations.[8][9]

Home range

[edit]

At night, or on hot or cloudy days, Epargyreus clarus rests on the underside of a leaf. It hangs upside down, holding its wings together over its back so that its silver spot is exposed. E. clarus is often restricted by its habitat, preferring to fly in the shade.[8]

Food resources

[edit]

Caterpillars

[edit]

Epargyreus clarus caterpillars consume leaves of herbs, vines, shrubs, and trees in the pea family (Fabaceae) including false indigobush (Amorpha fruticosa), American hogpeanut (Amphicarpaea bracteata), groundnut (Apios americana), American wisteria (Wisteria frutescens), Atlantic pigeonwings or butterfly pea (Clitoria mariana), and the introduced Dixie ticktrefoil (Desmodium tortuosum), kudzu (Pueraria montana), black locust (Robinia pseudoacacia), Chinese wisteria (Wisteria sinensis) and an assortment of other legumes.[9]

Innate host plant preferences confer greater performance on larvae, due to differences in leaf nutrient concentrations. However, preference may be influenced by a larva’s rearing host species.[10]

They live in meadows and swamps. Their average size is 2 inches (5 cm) long. Average life span is about 2 months. Their predators are wasps, ants, birds, frogs and spiders.[11]

Adults

[edit]

Adults use their long tongues to feed on the nectar of a variety of flowers, mud, and animal feces.[5] They almost never visit yellow flowers, favoring those that are blue, red, pink, purple, and sometimes white and cream. These include everlasting pea, common milkweed, red clover, buttonbush, blazing star, and thistles.[3]

Parental care

[edit]

Oviposition

[edit]

Females lay a single egg on or occasionally near host plants. They have been reported to oviposit on the least snoutbean (Rynchosia minima).[12]

Life cycle

[edit]

Eggs

[edit]

The eggs are green, with red coloration on top.[5]

Caterpillar

[edit]
Larva

Larvae are yellow with transverse subdorsal black stripes. The first thoracic segment is black, has a brown prothoracic shield, and is significantly thinner than its adjacent organs – the head and second thoracic segment. The head is described as black or reddish brown with two large, prominent anterior orange spots, which mimic eyes. The three true legs and midabdominal false legs are all bright orange.[9][5]

Shelter building

[edit]

All larval instars of Epargyreus clarus build shelters on their host plant. They build about five shelters throughout larval growth and development in four distinct styles. These are constructed from instinct and are unique to larval size. The first, second, and third instars make a simple, invariant structure that requires two incisions in the leaf and silk to fold over the flap created. Shelters are typically built on the apical portion of the leaflet. One study of E. clarus larvae in the Washington D.C. area defined five steps of shelter building:[4]

  1. Site selection: Larvae traverse across the entire leaf, following major veins and swinging their heads to create a single silk trail.
  2. Silk template: A thick, visible mat of silk outlines the intended incisions and ultimately becomes the ceiling of the leaf house. The larvae keep their posterior end on the template at all times.
  3. First cut: The first cut follows the silk template on the perimeter nearest the leaf apex. The larvae often interrupt chewing cuts in order to deposit silk at the terminus. The first cut is completed in about 6 to 7 segments. At the end, the larva makes a U-turn motion so that its position relative to the leaf surface and template margins remains the same while facing the opposite direction.
  4. Second cut and notch: The second cut follows the silk template on the edge closest to the base of the leaflet and is completed in about 2 to 4 segments. The first segment curves inwards, is made in a single effort and is about 75% the length of the first cut. The next segment is adjacent to the initial segment and continues past the length of the first cut. The juncture between the two segments creates a notch.
  5. Folding and securing: The uncut area between the first and second incisions composes the hinge. The hinge length tends to correspond to larval body length. The larvae lay silk strands along this area, parallel to the cut termini. As the strands contract, the notch creates a pinch, and the leaf flap is pulled over the leaflet plane. The pinch then becomes the peak of the roof and the larva begins to make "guy-wires" that attach the edge of the flap to the leaflet surface.

Larvae take breaks of about 30 minutes between each cut and fold. It typically takes less than 21/2 hours to complete shelter construction. Larvae spend about 95% of their time resting on the ceiling of the shelter.[4]

Adult
the silver-spotted skipper nectaring

First instar shelters are typically fastened by only about 2 to 4 "guy-wires". In addition, older caterpillars occasionally live in a nest made of multiple leaves connected by silk, especially when using host plants with smaller leaflets. For example, first, second, and third instars can make shelters with one Robinia leaflet, but the fourth and fifth instars have to use 2–4 or 5 leaflets.[4]

Pupa

[edit]

Larvae of Epargyreus clarus pupate throughout the winter. The pupa is dark brown with black and white marks. There are three or more overlapping generations of hibernating pupa in Florida – fewer in the northern regions.[5]

Adult

[edit]

The adult wingspan ranges from 1.75 to 2.625 inches (4.5 to 6.7 cm). Each forewing is triangular and dark brown with a large yellow-orange medial patch. Male forewings are more pointed than those of females. The hindwings are also dark brown, with a large silver patch on the discal third of the ventral side.[9][5]

Migration

[edit]

Adults are diurnal and fly from early to midsummer.[9] They have one brood per year in the North and West, two in the East, and three to four in the Deep South.[3]

Enemies

[edit]

Predators

[edit]
Wheel bug assassin bug vs. silver-spotted skipper caterpillar

Common predators of Epargyreus clarus include the paper wasp foragers Polistes fuscatus and Polistes dominula, as well as Crematogaster opuntiae ants. Studies have found that shelters protect larvae from these predators in lab settings. Shelters delayed or prevented Crematogaster opuntiae colonies from detecting the larvae. This is likely because of the ants' limited senses, in conjunction with the inhibitory structure of the shelter: they are able to detect a motionless caterpillar only within approximately one body length, and this is further obstructed by the single small opening into the shelter. While wasps in the laboratory setting did spend more time on areas of the leaf damaged by feeding or silk deposition, the larval shelter prevented visual detection and posed a physical barrier. However, shelters did not protect against these predators in the field. Wasps need spend only a few seconds on a leaflet to successfully identify and remove the larva from its shelter. This suggests that shelter identification and larval extraction is a learned ability, perhaps developed through visual or olfactory cues.[6]

Other predators include the sphecid wasp Stictia carolina, which sometimes supplies its nests with silver-spotted skipper larvae. Additional predators also often are attracted to the chemical cues from frass, or insect excrement. Epargyreus clarus larvae use an anal comb to throw their frass up to 38 body lengths away from them. When disturbed, larvae regurgitate an unpleasant, greenish chemical in defense.[5]

Parasites

[edit]

E. clarus larvae are a natural host of Baculovirus. The advanced stages of this infection cause larvae to feed without resting. Later, they climb to higher and more exposed areas, where they are more conspicuous to predators. Predator attack leaves their infected remains to be absorbed by the surrounding vegetation, which will be consumed by new hosts.[5]

Mating

[edit]

Instead of receiving ejaculate, a female receives a large spermatophore from a male for each copulation. Thus, the number of spermatophores a female contains roughly equates to the number of copulations. Generally, when there are more than two spermatophores present in a female, each one’s placement and degree of erosion indicates its age relative to the other. Epargyreus clarus is particularly resistant to erosion. Once mated, females seem to reject males, at least for some time afterwards.[13]

Male defense of places likely to attract females

[edit]

Males perch on branches and tall weeds about 3.3 to 6.6 feet (1 to 2 m) above the ground, darting out when any insect passes in hopes of finding a receptive Epargyreus clarus female. When confronted by another conspecific, male movements are swift and acrobatic. This perching behavior is part of a common mate-location strategy in which males compete for places where females are more likely to occur.[7] Females do not engage in this behavior, but may have to fly considerable distances to find mates, depending on the proximity of male territory to their larval development site. The places "defended" by males are roughly the size of a small room and are most likely chosen based on vegetation and topography.[8]

Physiology

[edit]

Vision

[edit]

Epargyreus clarus possesses compound eyes that lack pigment in the iris region. Each ommatidia, or single optical unit, has its own unique visual field that spans about 2°. The small visual field can be due to crystalline tracts in the eye that restrict the light to reach the retina only through this path.[14]

Interaction with plants

[edit]
Silver-spotted skipper nectaring Liatris

Epargyreus clarus is known to be a nectar-thief species. An individual nectar thief is an animal that takes nectar from a flower but does not pollinate it while doing so. A species as a whole can be considered a thief species if it does not pollinate a species of flowers during more than 50% of its feedings. Research has found that while foraging on flowers, the butterfly tended to probe the innermost disk florets, which are the male organs, but not make contact with the outermost florets, which are the female organs.[15]

Pest of crop plants

[edit]

Epargyreus clarus is known to feed on various crop plants such as soybean and kidney bean. However, no large-scale control measures have been taken, as the species' pest activity is not too detrimental.[5]

References

[edit]
  1. ^ "NatureServe Explorer 2.0 Epargyreus clarus Silver-spotted Skipper". explorer.natureserve.org. Retrieved 1 October 2020.
  2. ^ Kaufman, Kenn; Eaton, Eric R. (2007). Kaufman Field Guide to Insects of North America. Houghton Mifflin Books. pp. 236–237. ISBN 978-0-618-15310-7.
  3. ^ a b c d "Silver-spotted skipper". Butterflies and Moths of North America. Archived from the original on 2009-01-16. Retrieved 2008-10-11.
  4. ^ a b c d Weiss, Martha R.; Lind, Eric M.; Jones, Meg T.; Long, Jeremy D.; Maupin, Jennifer L. (2003-07-01). "Uniformity of Leaf Shelter Construction by Larvae of Epargyreus clarus (Hesperiidae), the Silver-Spotted Skipper". Journal of Insect Behavior. 16 (4): 465–480. doi:10.1023/A:1027399122333. ISSN 0892-7553. S2CID 28669353.
  5. ^ a b c d e f g h i j Hall, Donald W. (2017). "Silver-Spotted Skipper Epargyreus clarus (Cramer)". The Institute of Food and Agricultural Sciences Extension.
  6. ^ a b Jones, Meg T.; Castellanos, Ignacio; Weiss, Martha R. (2002-12-01). "Do leaf shelters always protect caterpillars from invertebrate predators?". Ecological Entomology. 27 (6): 753–757. Bibcode:2002EcoEn..27..753J. doi:10.1046/j.1365-2311.2002.00465.x. ISSN 1365-2311. S2CID 86245610.
  7. ^ a b Scott, James A. (1974). "Mate-Locating Behavior of Butterflies". The American Midland Naturalist. 91 (1): 103–117. doi:10.2307/2424514. JSTOR 2424514.
  8. ^ a b c Shapiro, Arthur M. (1978). "Phenotypic and Behavioral Convergence of "Silver-Spotted Skippers" (Lepidoptera: Hesperiidae)". Biotropica. 10 (2): 159–160. Bibcode:1978Biotr..10..159S. doi:10.2307/2388021. JSTOR 2388021.
  9. ^ a b c d e Miller, Jeffrey C.; Hammond, Paul C. (2003). LEPIDOPTERA OF THE PACIFIC NORTHWEST: CATERPILLARS AND ADULTS. p. 28.
  10. ^ Rosenwald, Laura C.; Lill, John T.; Lind, Eric M.; Weiss, Martha R. (2017-05-10). "Dynamics of host plant selection and host-switching by silver-spotted skipper caterpillars". Arthropod-Plant Interactions. 11 (6): 833–842. Bibcode:2017APInt..11..833R. doi:10.1007/s11829-017-9538-0. ISSN 1872-8855. S2CID 37474805.
  11. ^ Melissa Stewart, National Geographic Kids, Ick!, Washington DC, Text Copyright 2020
  12. ^ Scott, JA (1986). The Butterflies of North America: A Natural History and Field Guide. Stanford University Press. p. 583.
  13. ^ Burns, John M. (1968). "Mating Frequency in Natural Populations of Skippers and Butterflies as Determined by Spermatophore Counts". Proceedings of the National Academy of Sciences of the United States of America. 61 (3): 852–859. Bibcode:1968PNAS...61..852B. doi:10.1073/pnas.61.3.852. JSTOR 58962. PMC 305406. PMID 16591715.
  14. ^ Doving, K. B. (1969). "Function of Insect Compound Eyes Containing Crystalline Tracts". The Journal of General Physiology. 54 (2): 250–267. CiteSeerX 10.1.1.282.9479. doi:10.1085/jgp.54.2.250. PMC 2225925. PMID 4894303.
  15. ^ Venables, Adrienne; Barrows, Edward (1985). "SKIPPERS: POLLINATORS OR NECTAR THIEVES?" (PDF). Journal of the Lepidopterists' Society. 39 (4): 299–312.
[edit]