Monarch (butterfly) 246472 225821107 2008-07-15T15:41:47Z 76.120.155.178 /* Reproduction */ {{Taxobox | name = Monarch | image=Monarch_In_May.jpg | image_width=250px | image_caption=Female | image2=Monarch Butterfly Danaus plexippus Male 2664px.jpg | image2_width=250px | image2_caption=Male | regnum = [[Animal]]ia | phylum = [[Arthropod]]a | classis = [[Insect]]a | ordo = [[Lepidoptera]] | superfamilia = [[Papilionoidea]] | familia = [[Nymphalidae]] | subfamilia =[[Danainae]] | tribus = [[Danaini]] | genus = ''[[Danaus (genus)|Danaus]]'' | genus_authority = [[Jan Krzysztof Kluk|Kluk]], 1780 | species = '''''D. plexippus''''' | binomial = ''Danaus plexippus'' | binomial_authority = ([[Carolus Linnaeus|Linnaeus]], 1758) | synonyms = ''Danaus archippus'' <small>([[Johan Christian Fabricius|Fabricius]], 1793)<ref>{{cite book |others=Royal Entomological Society of London Committee on Generic Nomenclature, Committee on Generic Nomenclature |title=The Generic Names of British Insects |origyear=1934 |url=http://books.google.com/books?id=hgo1AAAAMAAJ&dq=%22Papilio+archippus+Fab.%22 |accessdate=2008-06-04 |date=2007-05-23 |publisher=British Museum (Natural History). Dept. of Entomology |pages=20}}</ref></small><br> ''Danaus menippe'' <small>([[Jacob Hübner|Hübner]], 1816)<ref>{{cite book |last=Scudder |first=Samuel H. |authorlink=Samuel Hubbard Scudder |coauthors=William M. Davis, Charles W. Woodworth, Leland O. Howard, Charles V. Riley, Samuel W. Williston |title=The butterflies of the eastern United States and Canada with special reference to New England |origyear=1989 |url=http://books.google.com/books?id=MOXnYGtuxx0C&dq=The+butterflies+of+the+eastern+United+States+and+Canada+with+special&q=menippe&pgis=1#search |accessdate=2008-06-04 |publisher=The author |pages=721 }}</ref></small><br> }} The '''monarch''' (''Danaus plexippus'') is a [[milkweed butterfly]] ([[subfamily]] Danainae), in the family [[Nymphalidae]]. It is perhaps the best known of all butterflies. Since the 19th century, it is also found in [[New Zealand]], and has been known in [[Australia]] since 1871. [http://www.amonline.net.au/factsheets/monarch.htm] [http://calamvalecreek.awardspace.com/regal.html] In Australia it is also known as the wanderer butterfly. [http://bugs.bio.usyd.edu.au/Entomology/internalAnatomy/imagePages/wandererCaterpillar.html] In Europe it is resident in the [[Canary Islands]] and [[Madeira]], and is found as a migrant in [[Russia]], [[Azores]], [[Sweden]] and [[Spain]]. Its wings feature an easily recognizable orange and black pattern, with a wingspan of {{nowrap|8.9&ndash;10.2 centimetres}} {{nowrap|(3½&ndash;4 in)}}.<ref name = Garber/> (The [[viceroy butterfly]] has a similar size, color, and pattern, but can be distinguished by an extra black stripe across the hindwing.) Female monarchs have darker veins on their wings, and the males have a spot in the center of each hindwing [http://www.dbc.uci.edu/~pjbryant/biodiv/lepidopt/danaidae/monarchm.htm] from which [[pheromone]]s are released. Males are also slightly larger. ==Taxonomy== The common name “monarch” was first published in 1874 by [[Samuel Hubbard Scudder|Samuel H. Scudder]] because “it is one of the largest of our butterflies, and rules a vast domain”.<ref name =Crispin>{{cite book |last=Guppy |first=Crispin S. |coauthors=Jon H. Shepard |title= Butterflies of British Columbia |origdate=2001-03-15 |url= http://books.google.ca/books?id=E-reg6vdutcC&pg=PA353&lpg=PA353&dq=Aegytus+Danaus+in+Greek+mythology+(Bird+et+al.+1995)&source=web&ots=b1uTTzyKr_&sig=IjljN2OVO20JRKaPeVfICO15_fU&hl=en#PPA353,M1 |accessdate=2008-03-29 |publisher=[[ University of British Columbia Press|UBC press]] |isbn= 0774808098 |pages=352–354}}</ref> But the name may be in honour of King [[William, Prince of Orange]].<ref>{{cite book |last=Jean |first=Adams |title=Insect Potpourri: Adventures in Entomology |url=http://books.google.ca/books?hl=en&lr=&id=KcqQ1rPZyRoC&oi=fnd&pg=PA28&dq=danaus+plexippus+wingspan&ots=yE1dSSQjCr&sig=54JduS0Zfn6CSbWc65l6QJ56bhE#PPA28,M1 |accessdate=2008-05-21 |year=1992 |publisher=CRC Press |isbn=1877743097 |pages=28-29}}</ref> The monarch was one of the many species originally described by [[Carolus Linnaeus|Linnaeus]] in his ''[[Systema Naturae]]'' of 1758. It was first placed in the genus ''[[Papilio]]''.<ref>{{cite book | last=Linnaeus | first=Carolus | authorlink=Carolus Linnaeus | title=Systema Naturae | location=Stockholm | publisher=Holmiae (Laurentii Salvii) | date=1758 | pages=p.471 | url=http://gdz.sub.uni-goettingen.de/no_cache/dms/load/img/?search=1&IDDOC=265138&pid=PPN362053006&page=471&zoomfactor=100&firstIDDOC=265101&lastIDDOC=265150&prevIDDOC=265137&nextIDDOC=265139&pageturn=475& | language=Latin | oclc=174638949 | accessdate=2008-02-04}}</ref> In 1780, [[Jan Krzysztof Kluk]] used the monarch as the [[type species]] for a new genus; ''[[Danaus (genus)|Danaus]]''. Since [[Latin grammar]] requires that the [[specific epithet]] and gender names agree it is unclear if the genus ''Danaus'' is drawn from [[Danaus]] (Greek [[wikt:Δαναός|Δαναός]]), a mythical king of Egypt and great-grandson of [[Zeus]] or is a masculinised version of [[Danaë]] (Greek [[wikt:Δανάη| Δανάη]]), Danaus’ great-great-granddaughter. The species name, ''plexippus'', refers to [[Plexippus]], one of the 50 sons of [[Aegyptus]], Danaus’ twin brother.<ref name =Crispin/><ref>{{cite book |last=Pyle |first=Robert Michael |title=Chasing Monarchs: Migrating with the Butterflies of Passage |url=http://books.google.com/books?id=gO9eXQI0ZxQC&pg=PA148&dq=monarch+danaus+mythology+danae&sig=VpblzAq4BdMap-RWe-AQWSA6raQ |accessdate=2008-05-14 |year=2001 |publisher=Houghton Mifflin Books |isbn=0618127437 |pages=148-149 }}</ref> [[Image:Monarch Butterfly Danaus plexippus on Echinacea purpurea 2800px.jpg|right|thumb|Underside of a ''Danaus plexippus plexippus'' feeding on nectar from an ''Echinacea purpurea'' flower]] The monarch is closely related to two very similar [[species]] which formed the ''Danaus (Danaus)'' [[subgenus]] before 2005. The first is the [[Jamaican monarch]] (''D. cleophile'') from [[Jamaica]] and [[Hispaniola]]. The second is the [[southern monarch]] (''D. erippus''), of [[South America]] south of the Amazon river. The southern monarch is almost indistinguishable from the monarch as an adult, the pupae are somewhat more different, and is often considered a subspeciesof the monarch proper. But analysis of [[Morphology (biology)|morphological]], [[mtDNA]] 12S [[rRNA]], [[Cytochrome c oxidase|cytochrome ''c'' oxidase]] [[subunit]] I, [[nuclear DNA]] 18S rRNA and [[Elongation factor|EF1]] subunit α [[DNA sequence|sequence]] data by Smith ''et al.'' (2005) indicates that it is better considered a distinct species. The separation of the monarch and southern monarch is comparatively recent. In all likelihood, the ancestors of the southern monarch separated from the monarch's population some 2 [[mya (unit)|mya]], at the end of the [[Pliocene]]. At the time [[sea level]]s were higher and the entire [[Amazonas]] lowland was a vast expanse of [[brackish]] swamp that offered hardly any butterfly habitat.<ref name = smith>{{cite journal |last=Smith |first=David A. |coauthors=Gugs Lushai and John A. Allen |year=2005 |month=June |title=A classification of ''Danaus'' butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA |journal=Zoological Journal of the Linnean Society |volume=144 |issue=2 |pages=191–212 |doi=10.1111/j.1096-3642.2005.00169.x |url=http://www.blackwell-synergy.com/action/showPdf?submitPDF=Full+Text+PDF+%28214+KB%29&doi=10.1111%2Fj.1096-3642.2005.00169.x |format=PDF |accessdate=2008-05-19}}</ref> Following the review of Smith ''et al.'' (2005), two sub-species of the monarch are recognized:<ref name = smith/> * ''Danaus p. plexippus'', the nominate subpecies, described by Linnaeus in 1758. It is the migratory subspecies known from most of North America. * ''Danaus p. megalippe'', named in 1826 by [[Jacob Hübner]]. It is the non-migratory subspecies, and is found from [[Florida]] and [[Georgia (U.S. state)|Georgia]] southwards, throughout the [[Caribbean]] and [[Central America]] to the [[Amazon River]]. Three local [[form (zoology)|form]]s were at first considered to be other subspecies, but are actually colour varieties of ''D. p. megalippe'': ** ''D. p. m.'' forma ''leucogyne'', named by [[Arthur Gardiner Butler|Arthur G. Butler]] in 1884. ** ''D. p. m.'' forma ''portoricensis'', named in 1941 by A.H. Clark. ** ''D. p. m.'' forma ''tobagi'', also named in 1941 by A.H. Clark. == Description == The monarch’s wingspan ranges from {{nowrap|8.9&ndash;10.2 centimetres}} {{nowrap|(3½&ndash;4 in)}}.<ref name = Garber>{{cite book |last=Garber |first=Steven D. |title=The Urban Naturalist |url=http://books.google.com/books?id=3w_X0A_Kq0gC&pg=PA76&dq=Danaus+plexippus+wingspan&lr=&sig=VAZ-Ig5m8stSDztbgZLOUH3-n3o#PPA79,M1 |accessdate=2008-05-26 |year=1998 |publisher=Courier Dover Publications |isbn=0486403998 |pages=76-79}}</ref> The upper side of the [[Insect wing|wings]] is tawny-orange, the veins and margins are black, and in the margins are two series of small white spots. The fore wings also have a few orange spots near the tip. The underside is similar but the tip of the forewing and hindwing are yellow-brown instead of tawny-orange and the white spots are larger.<ref name=Braby>{{cite book |last=Braby |first=Michael F. |title=Butterflies of Australia: Their Identification, Biology and Distribution |url=http://books.google.com/books?id=_GjN8R9V-nsC&pg=PA597&dq=Danaus+plexippus+wingspan&sig=ujr5eusQZHA2zIlpKQhPIcDjrpA#PPA598,M1 |accessdate=2008-05-26 |year=2000 |publisher=CSIRO Publishing |isbn=0643065911 |pages=597-599}}</ref> The male has a black patch of sex-scales on the hindwings, and the black veins on its wing are narrower than the female’s. The male is also slightly larger.<ref name=Braby/> The eggs are creamy white and later turn pale yellow. They are elongate and subconical, with approximately 23 longitudinal ridges and many fine traverse lines.<ref name=Braby/> A single egg weighs about {{nowrap|0.46 milligrams}}, and measures about {{nowrap|1.2 millimetres high}} and {{nowrap|0.9 millimetres wide}}.<ref>{{cite book |last=Oberhauser |first=Karen S. |coauthors=Michelle J. Solensky |title=The Monarch Butterfly: Biology and Conservation |url=http://books.google.com/books?id=H5nRqhCGA4UC&pg=RA1-PA21&dq=monarch+egg+size&sig=kbUhmh48vdX6vhiZZuanS89Lj7c#PRA1-PA3,M1 |accessdate=2008-06-03 |year=2004 |publisher=Cornell University Press |isbn=0801441889 |pages=3 }}</ref><ref>{{cite web |url=http://www.monarchwatch.org/biology/cycle1.htm |title=Monarch Life Cycle |accessdate=2008-06-06 |work=Biology |publisher=[http://www.monarchwatch.org Monarch Watch] }}</ref> == Migration == [[Image:Monarch butterflies in Santa Cruz 10.jpg|thumb|300px]left|Wintering monarchs in Santa Cruz [[California]] .]] Monarchs are especially noted for their lengthy annual migration. In North America they make massive southward migrations starting in August until the first frost. A northward migration takes place in the spring. (In Australia, they make limited migrations in cooler areas, [http://www.amonline.net.au/factsheets/monarch.htm] but the [[blue tiger butterfly]] is better known in Australia for its lengthy migration. [http://calamvalecreek.awardspace.com/regal.html]) Female monarchs deposit eggs for the next generation during these migrations. By the end of October, the population of the [[Rocky Mountains]] migrates to the [[sanctuary|sanctuaries]] of the [[Mariposa Monarca Biosphere Reserve]] in the [[Mexican state]]s of [[Michoacán]] and [[México (state)|México]]. The western population [[wikt:overwinter|overwinters]] in various sites in central coastal and southern [[California]], [[United States]], notably in [[Pacific Grove, California|Pacific Grove]] and [[Santa Cruz, California|Santa Cruz]]. The length of these journeys exceeds the normal lifespan of most monarchs, which is less than two months for butterflies born in early summer. The last generation of the summer enters into a non-reproductive phase known as diapause and may live seven months or more. [http://www.dbc.uci.edu/~pjbryant/biodiv/lepidopt/danaidae/monarchm.htm] During diapause, butterflies fly to one of many overwintering sites. The generation that overwinters generally does not reproduce until it leaves the overwintering site sometime in February and March. It is thought that the overwinter population of those east of the Rockies may reach as far north as [[Texas]] and [[Oklahoma]] during the spring migration. It is the second, third and fourth generations that return to their northern locations in the United States and [[Canada]] in the spring. How the species manages to return to the same overwintering spots over a gap of several generations is still a subject of research; the flight patterns appear to be inherited, based on a combination of [[circadian rhythm]] and the position of the sun in the sky.<ref> Gugliotta, Guy (2003): [http://ipm.osu.edu/trans/053_231.htm Butterflies Guided By Body Clocks, Sun Scientists Shine Light on Monarchs' Pilgrimage]. ''[[Washington Post]]'', May 23, 2003, page A03. Retrieved 2006-JAN-07.</ref> Monarch butterflies are one of the few insects capable of making transatlantic crossings. They are becoming more common in [[Bermuda]] due to increased usage of [[milkweed]] as an ornamental plant in flower gardens. Monarch butterflies born in Bermuda remain year round due to the island's mild climate. A few monarchs turn up in the far southwest of [[Great Britain]] in years when the wind conditions are right, and have been sighted as far east as Long Bennington. Monarchs can also be found in [[New Zealand]]. On the islands of [[Hawaii]] no migrations have been noted. == Reproduction == [[Image:Monarch Butterfly Danaus plexippus Vertical Caterpillar 2000px.jpg|right|thumb|Mature caterpillar feeding on [[swamp milkweed]]]] The mating period for the overwinter population occurs in the spring, just prior to migration from the overwintering sites. The courtship is fairly simple and less dependent on chemical pheromones in comparison with other species in its genus. [http://animaldiversity.ummz.umich.edu/site/accounts/information/Danaus_plexippus.html] Courtship is composed of two distinct stages, the aerial phase and the ground phase. During the aerial phase, the male pursues, nudges, and eventually takes down the female. Copulation occurs during the ground phase and involves the transfer of a spermatophore from the male to the female. Along with sperm, the spermatophore is thought to provide the female with energy resources that aid her in carrying out reproduction and remigration. The overwinter population returns only as far north as they need to go to find the early milkweed growth; in the case of the eastern butterflies that is commonly southern Texas. The life cycle of a monarch includes a change of form called complete [[metamorphosis (biology)|metamorphosis]]. The monarch goes through four radically different stages: # The eggs are laid by the females during spring and summer breeding months. # The eggs hatch (after 4 days), revealing worm-like larvae, the [[caterpillar]]s. The caterpillars consume their egg cases, then feed on [[milkweed]], and sequester substances called [[cardenolide]]s, a type of [[cardiac glycoside]]. During the caterpillar stage, monarchs store energy in the form of [[fat]] and [[nutrient]]s to carry them through the non-feeding [[pupa]] stage. # In the [[pupa]] or [[Pupa#Chrysalis|chrysalis]] stage, the caterpillar spins a silk pad on a twig, leaf, etc., and hangs from this pad by its last pair of prolegs. It hangs upside down in the shape of a 'J', and then molts, leaving itself encased in an articulated green [[exoskeleton]]. At this point, hormonal changes occur, leading to the development of a butterfly (metamorphosis). The chrysalis darkens (actually becomes transparent) a day before it emerges, and its orange and black wings can be seen. # The mature butterfly emerges after about two pupal weeks and hangs from the split chrysalis for several hours until its wings are dry (often in the morning). Meanwhile fluids are pumped into the crinkled wings until they become full and stiff. Some of this orangey fluid (called meconium) drips from the wings. Finally (usually in the afternoon) the monarch spreads its wings, quivers them to be sure they are stiff, and then flies away, to feed on a variety of flowers, including milkweed flowers, red clover, and goldenrod. Monarchs can live a life of two to eight weeks in a garden having their host ''[[Asclepias]]'' plants and sufficient flowers for nectar. This is especially true if the flower garden happens to be surrounded by native forest that seems to be lacking in flowers. ===Pictorial lifecycle=== <gallery> Image:Monarch Butterfly Showy Male 3000px.jpg|Monarch male showing its wings to attract a mate Image:Monarch Butterfly Danaus plexippus Mating Vertical 1800px.jpg|Monarch butterflies mating Image:Monarch Butterfly Danaus plexippus Laying Egg 2600px.jpg|Monarch butterfly laying eggs on [[Asclepias curassavica]] Image:Monarch Butterfly Eggs on Swan Plant.JPG|Monarch eggs on milkweed Image:Monarch caterpillar and egg.jpg|An early [[instar]] monarch caterpillar Image:Monarch Butterfly Danaus plexippus Feeding Down 3008px.jpg|A late instar caterpillar feeding Image:Danaus plexippus chrysalis 2003-10-16.jpg|Monarch butterfly chrysalis Image:Monarch Butterfly Emerging.JPG|Butterfly emerging from chrysalis Image:Monarch Butterfly Pink Zinnia 1800px.jpg|Adult monarch butterfly feeding on a ''[[Zinnia]]'' Image:Monarch butterflies in Santa Cruz-11.jpg‎|Wintering monarch butterflies </gallery> == Defense against predators == [[Image:Swamp Milkweed Asclepias incarnata Flowers Closeup 2800px.jpg‎|right|thumb|[[Swamp milkweed]], one of many species of ''[[Asclepias]]'' milkweeds used by the monarch]] Monarchs are foul-tasting and poisonous due to the presence of [[cardenolide aglycones]] in their bodies, which the caterpillars ingest as they feed on milkweed. [http://animaldiversity.ummz.umich.edu/site/accounts/information/Danaus_plexippus.html] Both forms advertise their unpalatability with bright colors and areas of high contrast on the skin or wings. This phenomenon is known as [[aposematism]]. Monarchs share this defense with the even more unpleasant-tasting and similar-appearing [[viceroy butterfly]], in an example of [[Müllerian mimicry]]. (Viceroys were at one time believed to be [[Mimicry#Batesian|Batesian]] mimics of monarchs.) [[Image:Monarch Viceroy Mimicry Comparison.jpg|550px|left|thumb|The monarch (left) and viceroy (right) butterflies exhibiting Müllerian mimicry]] {{clear}} ==Relationship with humans== [[Image:Camera november 091.jpg‎|right|thumb|A monarch butterfly in a butterfly garden]] The monarch is the [[List of U.S. state insects|state insect]] of Alabama,<ref>{{cite web |url=http://www.archives.state.al.us/emblems/st_insec.html |title=Official Alabama Insect |accessdate=2007-03-19 |date=[[2001-07-12]] |work=Alabama Emblems, Symbols and Honors |publisher=Alabama Department of Archives & History}}</ref> Idaho,<ref>{{cite web |url=http://www.shgresources.com/id/symbols/insect/ |title=Idaho Symbols, Insect: Monarch Butterfly |accessdate=2008-03-26 |work=Idaho State Symbols, Emblems, and Mascots |publisher=SHG resources, state handbook & guide}}</ref> Illinois,<ref>{{cite web |url=http://www.museum.state.il.us/exhibits/symbols/insect.html |title=State Symbol: Illinois Official Insect — Monarch Butterfly (''Danaus plexippus'') |accessdate=2008-03-26 |work=[Illinois] State Symbols |publisher=Illinois State Museum}}</ref> Minnesota,<ref>{{cite web |url= http://www.house.leg.state.mn.us/hinfo/govser/GOVSER2.pdf |format=PDF |accessdate=2008-03-26 |title=Minnesota State Symbols |publisher= Minnesota House of Representatives}}</ref> Texas,<ref>{{cite web |url=http://www.tsl.state.tx.us/ref/abouttx/symbols.html |accessdate=2008-03-26 |title=Texas State Symbols |publisher=The Texas State Library and Archives}}</ref> and the [[List of U.S. state butterflies|state butterfly]] of Vermont<ref>{{cite web |url=http://www.libraries.vermont.gov/www/html/emblems/butterfly.htm |accessdate=2008-03-26 |title=(Vermont) State Butterfly |publisher=Vermont Department of Libraries}}</ref> and West Virginia.<ref>{{cite web |url=http://www.wv.gov/govdirectory/statgeninfo.pdf |accessdate=2008-03-26 |format=PDF |title=West Virginia Statistical Information, General State Information |publisher=Official West Virginia Web Portal}}</ref> It was nominated in 1990 as the national insect of the [[United States|United States of America]], along with the honeybee (''[[Apis mellifera]]''),<ref>{{cite news |first=Nicholas |last=Wade |title=Choosing a National Bug |publisher=[[The New York Times]] |date=1990-06-01 |accessdate=2008-03-26 |url=http://query.nytimes.com/gst/fullpage.html?res=9C0CE4D81439F932A35755C0A966958260}}</ref> but the legislation did not pass.<ref>{{cite web |url=http://www.entsoc.org/resources/faq.htm#triv7 |accessdate=2008-03-26 |title=Frequently Asked Questions on Entomology |publisher=Entomological Society of America}}</ref> Many people like to attract monarchs by growing a [[Butterfly gardening|butterfly garden]]. Others enjoy raising them for pleasure or for educational purposes. For migrating flocks, sanctuaries have been created at favorite wintering locations.<ref>{{cite web |url= http://www.sciencedaily.com/videos/2005/0810-saving_butterflies.htm |title= Saving Butterflies Insect Ecologist Spearheads Creation of Oases for Endangered Butterflies |accessdate=2008-05-27 |publisher= ScienceDaily |date=2005-01-01}}</ref> Even [[tourism]] revenue is thus generated. Many schools also enjoy growing, and attending to monarch butterflies, starting with the caterpillar form. When the butterflies reach adulthood they are then released into the wild.<ref>{{cite web |url= http://www.monarchwatch.org/ |title= Monarch Watch |accessdate=2008-05-27 |publisher= The Kansas Biological Survey, Univ of Kansas |date= 2008-05-27 }}</ref> ==Threats== Recent illegal deforestation of the monarch's overwintering grounds have led to a drastic reduction in the butterfly's population. Efforts to classify it as a protected species and to restore its habitat are under way. Mexican authorities expected a significant increase in the monarch population in the 2005-2006 season. ===Predators=== Although monarchs feed on milkweed, variations in the quantity of cardiac glycosides exist between species, individuals and even parts of the host plant. The levels of toxins in adult monarchs reflect the levels in their host plants. This means some monarchs are not foul-tasting but are [[Batesian mimicry|Batesian]] or [[Mimicry#Automimicry|auto-mimics]]. Some species of predators have learnt to measure the toxins by taste and reject butterflies with high cardiac glycosides contents, eating only the ones with low cardiac glycosides contents. In the butterfly, the cardiac glycosides are concentrated in the abdomen and wings. Some species of predators differentiate these parts and consume only the most palatable ones.<ref name= Barbosa>{{cite book |last=Barbosa |first=Pedro |coauthors= Deborah Kay Letourneau |title=Novel Aspects of Insect-plant Interactions |url= http://books.google.fr/books?id=2fZbhllItE4C&pg=PA45&dq=monarch+predators+asclepias&sig=bnjfIdmBqSFoBcB9L6FPAVoeTTA#PPA31,M1 |accessdate=2008-05-12 |year=1988 |publisher=Wiley-Interscience |isbn= 0471832766 |pages=29-31 |chapter=5 }}</ref> Bird predators include [[Brown Thrasher]]s, [[Grackle]]s, [[American Robin|Robin]]s, [[Cardinal (bird)|Cardinal]]s, [[Sparrow]]s, [[Scrub Jay]]s and [[Pinyon Jay]]s.<ref name= Barbosa/> In North America, eggs and first instar larvae of the monarch are eaten by larvae and adults of the [[Introduced species|introduced]] Asian lady beetle (''[[Harmonia axyridis]]'').<ref>{{cite journal |last=Koch |first=R. L. |coauthors=W. D. Hutchison, R. C. Venette and G. E. Heimpel |year=2003 |month=October |title=Susceptibility of immature monarch butterfly, ''Danaus plexippus'' (Lepidoptera: Nymphalidae: Danainae), to predation by ''Harmonia axyridis'' (Coleoptera: Coccinellidae) |journal=Biological Control |volume=28 |issue=2 |pages=265–270 |doi=10.1016/S1049-9644(03)00102-6 |url=http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WBP-48TKDRR-6&_user=1072191&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000051289&_version=1&_urlVersion=0&_userid=1072191&md5=fb8ace3d840cb24a87d5d7276ef7bb6c |accessdate=2008-04-23 }}</ref> [[Image:albino monarch butterfly.jpg|right|thumb|White [[Polymorphism (biology)|morph]] of the monarch in Hawaii]] On [[Oahu]], a white [[Polymorphism (biology)|morph]] of the monarch has emerged. This is because of the [[Introduced species|introduction]], in 1965 and 1966, of two [[bulbul]] species, ''[[Pycnonotus cafer]]'' and ''Pycnonotus jacosus''. They are now the most common insectivore birds, and probably the only ones preying on insects as big as the monarch. Monarchs in [[Hawaii]] are known to have low cardiac glycoside levels, but the birds may also be [[Physiological tolerance|tolerant]] for the chemical. The two species hunt the larvae and some pupae from the branches and underside of leaves in milkweed bushes. The bulbuls also eat resting and ovipositing adults, but rarely flying ones. Because of its colour the white morph has a higher survival rate than the orange one. This is either because of [[apostatic selection]] (i.e. the birds have learnt the orange monarchs can be eaten) or because of [[camouflage]]; the white morph matches the white pubescence of milkweed, or the patches of light shining through foliage.<ref>{{cite journal |last=Stimson |first=John |coauthors=Mark Berman |year=1990 |title= Predator induced colour polymorphism in Danaus plexippus L. (Lepidoptera: Nymphalidae) in Hawaii |journal=Heredity |volume=65 |issue=3 |pages=401–406 |accessdate=2008-04-06 |laysummary=http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=2472199&q=predator+induced+colour+polymorphism&uid=792450984&setcookie=yes |doi= 10.1038/hdy.1990.110 }}</ref> ===Parasites=== Parasites include the [[Tachinidae|tachinid]] fly ''Lesperia archippivora''. Parasitized larvae complete their moult, suspend, but die before pupation. At that time one white [[maggot]] comes out of the larvae, suspended by a [[silk|silken]] thread. The maggot then forms a brown pupa on the ground.<ref name = Brewer>{{cite journal |last=Brewer |first=Jo |coauthors=Gerard M. Thomas |year=1966 |title=Causes of death encountered during rearing of Danaus plexippus (Danaidae) |journal=Journal of The Lepidopterist’s Society |volume=20 |issue=4 |pages=235–238 |url=http://research.yale.edu/peabody/jls/pdfs/1960s/1966/1966-20(4)235-Brewer.pdf |format=PDF |accessdate= 2008-04-13 | laysummary =http://research.yale.edu/peabody/jls/htms/1960s/1966-20(4)235-Brewer.htm }}</ref> The bacteria ''[[Micrococcus]] flacidifex danai'' also infects the larvae and causes “black death”. As usual, just before pupation the larvae migrate to a horizontal surface. They die a few hours later, attached only by one pair of prolegs, thorax and abdomen hanging limp. The body turns black shortly after. The bacteria ''[[Pseudomonas]] aeriginosa'' has no invasive powers, but causes [[secondary infection]]s in weakened insects. It is a common cause of death in laboratory reared insects.<ref name = Brewer/> The [[protozoa]]n ''[[Ophryocystis elektroscirrha]]'' is another parasite of the monarch. It infects the [[hypodermis|subcutaneous tissues]] and propagates by [[spore]]s formed during the pupal stage. The spores are found over all of the body of infected butterflies, with the greatest number on the abdomen. These spores are passed, from [[vertical transmission|female to caterpillar]], when spores rub off during egg-laying and are then ingested by caterpillars. Severely infected individuals are weak, unable to expand their wings, or unable to eclose, and have a shortened lifespan but probably occur a low frequencies in nature. This is not the case in laboratory or commercial rearing, where after a few generations all individuals can be infected.<ref>{{cite journal |last=Leong |first= K. L. H. |coauthors=M. A. Yoshimura, H. K. Kaya and H. Williams |year=1997 |title= Instar Susceptibility of the Monarch Butterfly (''Danaus plexippus'') to the Neogregarine Parasite, ''Ophryocystis elektroscirrha'' |journal=Journal of Invertebrate Pathology |volume=69 |issue=1 |pages=79–83 |accessdate=2008-04-15 |doi=10.1006/jipa.1996.4634 |laysummary=http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WJV-45S96SB-D&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=7d4874a42d109805f3a1e77129046e4f }}</ref> ==References== {{reflist|2}} {{Refimprove|date=December 2007}} ==External links== {{commons|Danaus plexippus}} {{Wikispecies|Danaus plexippus}} * [http://depts.washington.edu/natmap/facts/monarch_712.html Monarch butterfly facts and photos] - NatureMapping Program * [http://www.youtube.com/watch?v=l67SLCOYoHA#GU5U2spHI_4 Video of a monarch butterfly emerging from chrysalis] * [http://www.amonline.net.au/factsheets/monarch.htm Australian Museum fact sheet on monarch butterflies] * [http://calamvalecreek.awardspace.com/regal.html Critters of Calamvale Creek &mdash; a monarch tells its story] * [http://www.learner.org/jnorth/monarch/ Monarch Migration Maps] * [http://www.monarchwatch.org/ Monarch Watch] of the [[University of Kansas]] [[Entomology]] Department * [http://www.michoacanmonarchs.org/ Michoacan Reforestation Fund] * [http://www.butterfliesandmoths.org/species?l=1892 USGS description of monarch butterfly] * [http://www.monarchparasites.org/ MonarchHealth!] * [http://www.monarch.org.nz/ Monarch Butterfly Trust NZ] * [http://www.geocities.com/wyllz/id177 Monarch butterfly life cycle photographs] * [http://www.morethanmonarchs.org More Than Monarchs] * [http://www.pbase.com/10kzoomfz/wild_monarch_07proj_eggeclose Monarch egg eclosion] [[Category:Danaus]] [[Category:Butterflies and moths of North America]] [[Category:Pollinators]] [[Category:Fauna of the United States]] [[Category:Fauna of Mexico]] [[Category:Natural history of North America]] [[cs:Monarcha stěhovavý]] [[da:Monark (sommerfugl)]] [[de:Monarchfalter]] [[es:Danaus plexippus]] [[fr:Monarque (papillon)]] [[it:Danaus plexippus]] [[he:דנאית מלכותית]] [[hu:Pompás királylepke]] [[nl:Monarchvlinder]] [[ja:オオカバマダラ]] [[no:Monarksommerfugl]] [[pl:Danaus plexippus]] [[pt:Borboleta-monarca]] [[ro:Fluture Monarh]] [[fi:Monarkkiperhonen]] [[tl:Danaus plexippus]] [[zh:黑脉金斑蝶]]