Editing Butterfly (section)

===Ecology===

====Parasitoids, predators, and pathogens====

[[File:Braconid parasitoid wasp Apanteles sp eggs & Lime Butterfly (Papilio demoleus) cat W IMG 2862.jpg|thumb|[[Braconid]] [[parasitoid]]al wasp (''[[Apanteles]]'' species) cocoons attached to lime butterfly (''[[Papilio demoleus]]'') caterpillar]]

Butterflies are threatened in their early stages by [[parasitoid]]s and in all stages by predators, diseases and environmental factors. [[Braconidae|Braconid]] and other parasitic wasps lay their eggs in lepidopteran eggs or larvae and the wasps' parasitoid larvae devour their hosts, usually pupating inside or outside the desiccated husk. Most wasps are very specific about their host species and some have been used as biological controls of pest butterflies like the [[Pieris brassicae|large white butterfly]].<ref name=Feltwell401>{{cite book |author=Feltwell, J. |title=Large White Butterfly: The Biology, Biochemistry and Physiology of ''Pieris brassicae'' (Linnaeus) |url=https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA401 |year=2012 |publisher=Springer |isbn=978-94-009-8638-1 |pages=401– |access-date=8 January 2016 |archive-date=10 May 2016 |archive-url=https://web.archive.org/web/20160510071358/https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA401 |url-status=live }}</ref> When the [[Pieris rapae|small cabbage white]] was accidentally introduced to New Zealand, it had no natural enemies. In order to control it, some pupae that had been parasitised by a chalcid wasp were imported, and natural control was thus regained.<ref>{{cite book |author1=Burton, Maurice |author2=Burton, Robert |title=International Wildlife Encyclopedia: Brown bear - Cheetah |url=https://books.google.com/books?id=WLIOwLemzyQC&pg=PA416 |year=2002 |publisher=Marshall Cavendish |isbn=978-0-7614-7269-8 |page=416 |access-date=8 January 2016 |archive-date=9 May 2016 |archive-url=https://web.archive.org/web/20160509074556/https://books.google.com/books?id=WLIOwLemzyQC&pg=PA416 |url-status=live }}</ref> Some flies lay their eggs on the outside of caterpillars and the newly hatched fly larvae bore their way through the skin and feed in a similar way to the parasitoid wasp larvae.<ref name=Allen>{{cite book |author=Allen, Thomas J. |title=A Field Guide to Caterpillars |url=https://books.google.com/books?id=SS6s7tck0fIC&pg=PA15 |year=2005 |publisher=Oxford University Press |isbn=978-0-19-803413-1 |page=15 |access-date=8 January 2016 |archive-date=27 April 2016 |archive-url=https://web.archive.org/web/20160427045848/https://books.google.com/books?id=SS6s7tck0fIC&pg=PA15 |url-status=live }}</ref> Predators of butterflies include ants, spiders, wasps, and birds.<ref>{{cite web |title=Parasites and Natural Enemies |url=http://monarchlab.org/biology-and-research/biology-and-natural-history/parasites-natural-enemies |publisher=University of Minnesota |access-date=16 October 2015 |archive-date=7 October 2015 |archive-url=https://web.archive.org/web/20151007132714/http://monarchlab.org/biology-and-research/biology-and-natural-history/parasites-natural-enemies/ |url-status=live }}</ref>

Caterpillars are also affected by a range of bacterial, viral and fungal diseases, and only a small percentage of the butterfly eggs laid ever reach adulthood.<ref name=Allen/> The bacterium ''[[Bacillus thuringiensis]]'' has been used in sprays to reduce damage to crops by the caterpillars of the large white butterfly, and the [[entomopathogenic fungus]] ''[[Beauveria bassiana]]'' has proved effective for the same purpose.<ref name=Feltwell429>{{cite book |author=Feltwell, J. |title=Large White Butterfly: The Biology, Biochemistry and Physiology of ''Pieris Brassicae'' (Linnaeus) |url=https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA429 |year=2012 |publisher=Springer |isbn=978-94-009-8638-1 |page=429 |access-date=8 January 2016 |archive-date=5 May 2016 |archive-url=https://web.archive.org/web/20160505022425/https://books.google.com/books?id=8XnrCAAAQBAJ&pg=PA429 |url-status=live }}</ref>

====Endangered species====

[[Queen Alexandra's birdwing]], found in [[Papua New Guinea]], is the largest butterfly in the world. The species is [[List of endangered insects#Other Lepidoptera species|endangered]], and is one of only three insects (the other two being butterflies as well) to be listed on [[CITES#Appendix I|Appendix I]] of [[CITES]], making international trade illegal.<ref name="CITES appendices">[http://www.cites.org/eng/app/appendices.php CITES appendices I, II and III] {{Webarchive|url=https://web.archive.org/web/20171205014647/http://www.cites.org/eng/app/appendices.php |date=5 December 2017 }}, official website</ref>

====Defences====

{{further|Defense in insects|Anti-predator adaptation|Mimicry|Seasonal polyphenism}}

Butterflies protect themselves from predators by a variety of means.

Chemical defences are widespread and are mostly based on chemicals of plant origin. In many cases the plants themselves evolved these toxic substances as [[plant defense against herbivory|protection]] against herbivores. Butterflies have evolved mechanisms to sequester these plant toxins and use them instead in their own defence.<ref>{{cite journal |doi=10.1146/annurev.ento.47.091201.145121 |last1=Nishida |first1=Ritsuo |year=2002 |title=Sequestration of Defensive Substances from Plants by Lepidoptera |journal=Annual Review of Entomology |volume=47 |pages=57–92 |pmid=11729069}}</ref> These defence mechanisms are effective only if they are well advertised; this has led to the evolution of bright colours in unpalatable butterflies ([[aposematism]]). This signal is commonly [[mimicry|mimicked]] by other butterflies, usually only females. A [[Batesian mimicry|Batesian mimic]] imitates another species to enjoy the protection of that species' aposematism.<ref name=Edmunds/> The [[Papilio polytes|common Mormon]] of India has female morphs which imitate the unpalatable red-bodied swallowtails, the [[Pachliopta aristolochiae|common rose]] and the [[Pachliopta hector|crimson rose]].<ref>{{cite web |last1=Halloran |first1=Kathryn |last2=Wason |first2=Elizabeth |title=''Papilio polytes'' |url=http://animaldiversity.org/accounts/Papilio_polytes/ |website=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |access-date=12 September 2015 |date=2013 |archive-date=7 October 2015 |archive-url=https://web.archive.org/web/20151007182812/http://animaldiversity.org/accounts/Papilio_polytes/ |url-status=live }}</ref> [[Müllerian mimicry]] occurs when aposematic species evolve to resemble each other, presumably to reduce predator sampling rates; ''[[Heliconius]]'' butterflies from the Americas are a good example.<ref name=Edmunds>{{cite book |author=Edmunds, M. |year=1974 |title=Defence in Animals |url=https://archive.org/details/defenceinanimals0000edmu |url-access=registration |publisher=Longman |pages=[https://archive.org/details/defenceinanimals0000edmu/page/74 74]–78, 100–113}}</ref>

[[Camouflage]] is found in many butterflies. Some like the oakleaf butterfly and [[Doleschallia bisaltide|autumn leaf]] are remarkable imitations of leaves.<ref>{{cite journal |doi=10.1086/283868 |last1=Robbins |first1=Robert K. |s2cid=34146954 |year=1981 |title=The "False Head" Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies |journal=American Naturalist |volume=118 |issue=5| pages=770–775| bibcode=1981ANat..118..770R }}</ref> As caterpillars, many defend themselves by freezing and appearing like sticks or branches.<ref>{{cite book| author=Forbes, Peter |date=2009 |title=Dazzled and Deceived: Mimicry and Camouflage |publisher=Yale University Press |isbn=978-0-300-17896-8|title-link=Dazzled and Deceived: Mimicry and Camouflage }}</ref> Others have [[deimatic]] behaviours, such as rearing up and waving their front ends which are marked with eyespots as if they were snakes.<ref name=Springer>{{cite web |url=http://www.springerreference.com/docs/html/chapterdbid/86053.html |title=Deimatic Behavior |publisher=Springer |year=2012 |access-date=31 December 2012 |author=Edmunds, Malcolm |archive-date=28 July 2013 |archive-url=https://web.archive.org/web/20130728153247/http://www.springerreference.com/docs/html/chapterdbid/86053.html |url-status=live }}</ref> Some papilionid caterpillars such as the giant swallowtail (''[[Papilio cresphontes]]'') resemble bird droppings so as to be passed over by predators.<ref>{{cite web |title=Featured Creatures: Giant Swallowtail |url=http://entnemdept.ifas.ufl.edu/creatures/citrus/giantswallowtail.htm |publisher=University of Florida |access-date=12 September 2015 |archive-date=11 June 2019 |archive-url=https://web.archive.org/web/20190611201118/https://entnemdept.ifas.ufl.edu/creatures/citrus/giantswallowtail.htm |url-status=live }}</ref> Some caterpillars have hairs and bristly structures that provide protection while others are gregarious and form dense aggregations.<ref name=Edmunds/> Some species are [[myrmecophile]]s, forming [[Symbiosis|mutualistic associations]] with [[ant]]s and gaining their protection.<ref>{{cite journal|author1=Fiedler, K. |author2=Holldobler, B. |author3=Seufert, P. |title=Butterflies and Ants: The Communicative Domain |journal=Cellular and Molecular Life Sciences |volume=52 |date=1996 |pages=14–24 |doi=10.1007/bf01922410|s2cid=33081655 }}</ref> Behavioural defences include perching and angling the wings to reduce shadow and avoid being conspicuous. Some female [[Nymphalid]] butterflies guard their eggs from parasitoidal [[wasp]]s.<ref>{{cite journal |last1 = Nafus |first1 = D. M. |last2 = Schreiner |first2 = I. H. |year = 1988 |title = Parental Care in a Tropical Nymphalid Butterfly ''Hypolimas anomala'' |journal = Animal Behaviour |volume = 36 |issue = 5| pages = 1425–1443 |doi=10.1016/s0003-3472(88)80213-6| s2cid = 53183529 }}</ref>

The Lycaenidae have a false head consisting of eyespots and small tails (false antennae) to deflect attack from the more vital head region. These may also cause ambush predators such as spiders to approach from the wrong end, enabling the butterflies to detect attacks promptly.<ref>{{cite journal |author=Cooper, William E. Jr. |year=1998 |title=Conditions Favoring Anticipatory and Reactive Displays Deflecting Predatory Attack |journal=Behavioral Ecology |volume=9 |issue=6 |pages=598–604 |doi=10.1093/beheco/9.6.598|citeseerx=10.1.1.928.6688 }}</ref><ref>{{cite journal |doi=10.1017/S1464793105006810 |last1=Stevens |first1=M. |s2cid=24868603 |year=2005 |title=The Role of Eyespots as Anti-Predator Mechanisms, Principally Demonstrated in the Lepidoptera |journal=Biological Reviews |volume=80 |issue=4| pages=573–588 |pmid=16221330}}</ref> Many butterflies have [[Eyespot (mimicry)|eyespots]] on the wings; these too may deflect attacks, or may serve to attract mates.<ref name="pmid11676917" /><ref name ="Eyespot">{{cite journal |author=Brakefield, PM |year=1996 |title=Development, Plasticity and Evolution of Butterfly Eyespot Patterns |journal=Nature |issue=6606| pages=236–242 |volume=384 |doi=10.1038/384236a0 |pmid=12809139| last2=Gates |first2=Julie |last3=Keys |first3=Dave |last4=Kesbeke |first4=Fanja |last5=Wijngaarden |first5=Pieter J. |last6=Montelro |first6=Antónia |last7=French |first7=Vernon |last8=Carroll |first8=Sean B. |display-authors=etal| bibcode=1996Natur.384..236B |s2cid=3341270 }}</ref>

Auditory defences can also be used, which in the case of the [[grizzled skipper]] refers to vibrations generated by the butterfly upon expanding its wings in an attempt to communicate with ant predators.<ref>{{Cite journal|last=Elfferich|first=Nico W.|date=1998|title=Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants|url=http://natuurtijdschriften.nl/search?identifier=538588|journal=Deinsea|volume=4|issue=1|access-date=6 November 2017|archive-date=27 September 2017|archive-url=https://web.archive.org/web/20170927000340/http://natuurtijdschriften.nl/search?identifier=538588|url-status=live}}</ref>

Many tropical butterflies have [[seasonal polyphenism|seasonal forms]] for dry and wet seasons.<ref>{{cite journal |pmid=18811432 |date=December 1998 |author1=Brakefield, P. M. |author2=Kesbeke, F. |author3=Koch, P. B. |title=The Regulation of Phenotypic Plasticity of Eyespots in the Butterfly ''Bicyclus anynana'' |volume=152 |issue=6 |pages=853–60 |issn=0003-0147 |doi=10.1086/286213 |journal=The American Naturalist|bibcode=1998ANat..152..853B |s2cid=22355327 }}</ref><ref>{{cite journal |doi=10.1006/mpev.2000.0872 |last1=Monteiro |first1=A. |last2=Pierce |first2=N. E. |s2cid=20314608 |year=2001 |title=Phylogeny of Bicyclus (Lepidoptera: Nymphalidae) Inferred from COI, COII, and EF-1 Alpha Gene Sequences |journal=Molecular Phylogenetics and Evolution |volume=18 |issue=2 |pages=264–281 |pmid=11161761| bibcode=2001MolPE..18..264M |url=http://pdfs.semanticscholar.org/ab63/4db58a38ab7e6d3f77f71e96f16157b0b37b.pdf |archive-url=https://web.archive.org/web/20190303094544/http://pdfs.semanticscholar.org/ab63/4db58a38ab7e6d3f77f71e96f16157b0b37b.pdf |url-status=dead |archive-date=2019-03-03 }}</ref> These are switched by the hormone [[ecdysone]].<ref>{{cite journal |pmid=12492404 |date=Jan 2003 |author=Nijhout, Hf |s2cid=6027259 |title=Development and Evolution of Adaptive Polyphenisms |volume=5 |issue=1 |pages=9–18 |issn=1520-541X |journal=Evolution & Development |doi=10.1046/j.1525-142X.2003.03003.x}}</ref> The dry-season forms are usually more cryptic, perhaps offering better camouflage when vegetation is scarce. Dark colours in wet-season forms may help to absorb solar radiation.<ref>{{cite journal |last1=Brakefield |first1=Paul M. |last2=Larsen |first2=Torben B. |title=The Evolutionary Significance of Dry and Wet Season Forms in some Tropical Butterflies |year=1984 |journal=Biological Journal of the Linnean Society |volume=22 |doi=10.1111/j.1095-8312.1984.tb00795.x |pages=1–12 |hdl=1887/11011 |url=https://openaccess.leidenuniv.nl/bitstream/handle/1887/11011/029_009.pdf?sequence=1 |hdl-access=free |access-date=23 September 2019 |archive-date=27 July 2020 |archive-url=https://web.archive.org/web/20200727075120/https://openaccess.leidenuniv.nl/bitstream/handle/1887/11011/029_009.pdf?sequence=1 |url-status=live }}</ref><ref>{{cite journal |doi=10.1098/rspb.2003.2571 |last1=Lyytinen |first1=A. |last2=Brakefield |first2=P. M. |last3=Lindström |first3=L. |last4=Mappes |first4=J. |year=2004 |title=Does Predation Maintain Eyespot Plasticity in ''Bicyclus anynana'' |journal=Proceedings of the Royal Society B |volume=271 |issue=1536| pages=279–283| pmc=1691594 |pmid=15058439}}</ref><ref name ="Eyespot"/>

Butterflies without defences such as toxins or mimicry protect themselves through a flight that is more bumpy and unpredictable than in other species. It is assumed this behavior makes it more difficult for predators to catch them, and is caused by the [[turbulence]] created by the small whirlpools formed by the wings during flight.<ref>{{Cite web|url=https://www.insidescience.org/news/mathematical-butterfly-simulations-provide-new-insights-flight|title=The Mathematical Butterfly: Simulations Provide New Insights On Flight|website=Inside Science|date=19 April 2013 |access-date=15 May 2018|archive-date=16 May 2018|archive-url=https://web.archive.org/web/20180516014712/https://www.insidescience.org/news/mathematical-butterfly-simulations-provide-new-insights-flight|url-status=live}}</ref>

<gallery class=center mode=nolines widths="180px" heights="180px">
File:Heliconius mimicry.png|''[[Heliconius]]'' warns off predators with [[Müllerian mimicry]].<ref>{{cite journal |doi=10.1371/journal.pbio.0040341 |date=October 2006 |last=Meyer |first=A. |title=Repeating Patterns of Mimicry |volume=4 |issue=10 |pages=e341 |issn=1544-9173 |pmid=17048984 |pmc=1617347 |journal=PLOS Biology |doi-access=free }}</ref>
File:Papilio cresphontes larva defensive.JPG|[[Giant swallowtail]] caterpillar everting its [[osmeterium]] in defence; it is also [[mimesis (biology)|mimetic]], resembling a bird dropping.
File:Bird-damaged Speckled Wood Pararge aegeria.JPG|Eyespots of [[Speckled wood (butterfly)|speckled wood]] (''Pararge aegeria'') distract predators from attacking the head. This insect can still fly with a damaged left hindwing.
</gallery>