MARIPOSAS de BOLIVIA

Enero - Febrero 2012

first update d. 6 November 2015

Last update d. 24 April 2017

Redactor: Lars Andersen


Longwing, Heliconiini species (Swainson, 1822)

Enero - Febrero 2005 - 12 sitio 71

Coroico 1900 m.h. d. 12 February 2012. Photographer Lars AndersenHeliconius telesiphe on Heliotropium flowers. Coroico 1300 m.h. d. 14 february 2012. Photographer Lars AndersenHeliconius telesiphe. Yolosa, Yungas, elev. 1400 m. 19 January 2012. Photographer: Lars Andersen Philaethria dido. Coroico 1900 m.h. d. 17 january 2012. Photographer Lars Andersen

Introduction

The tribe Heliconiini, colloquially known as Longwings, includes 71 species, all confined exclusively to the neotropics (from southern Florida, Texas, Mexico, Central America pass through the Amazon to Paraguay and northern Argentina).

The Heliconiini includes the genera Heliconius, Podotricha, Dryas, Agraulis, Dione, Dryadula, Eueides, Neruda, Laparus and Philaethria.

All Heliconius species have elongated black wings, marked with simple but striking patterns usually featuring streaks or patches of red and cream, or blue and cream. A few, such as Heliconius sara, antiochus and wallacei have a metallic blue sheen over the basal area of both wings. All are characterised by their delicate fluttering flight, long straight antennae, and fondness for flowers.

The 39 Heliconius species are much studied by geneticists and taxonomists. Many of them produce a staggering variety of colour forms - Heliconius erato e.g. produces no less than 29 geographical forms, each of which corresponds almost exactly in colour and pattern to a 'sister' subspecies of Heliconius melpomene flying in the same area.

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Introduktion

Underfamilie Heliconiinae, famlien Nymphalidae, også kendt som Langvinger, omfatter 72 arter, alle geografisk udbredt i Neotroperne fra Florida, Texas, Mexico til Amazonas og det nordlige Argentina og Paraguay. Underfamilie Heliconiinae (som udover tribe Heliconiini, Langvinger også rummer tribe Argynnini, Perlemorsommerfugle med 120 arter udbredt overalt fra polar til tropisk klima undtagen på Antarktisk).

Langvinger, Heliconiini omfatter slægterne Heliconius, Podotricha, Dryas, Agraulis, Dione, Dryadula, Eueides, Neruda, Laparus og Philaethria.

Alle Heliconius arter har aflange sorte vinger, mærket med enkle, iøjnefaldende mønstre med striber eller pletter af rød og lysegul, eller blå og creme. Enkelte arter som Heliconius sara, H. antiochos og H. wallacei har metallisk blå skær over det basale område på begge vinger.

Alle Langvinger er kendetegnet ved deres sarte flagrende flugt, lange lige antenner, og forkærlighed til blomster.

De 39 Heliconius arter er blandt genetikere og taksonomer de mest undersøgte sommerfugle fra troperne. Mange af arter findes i en svimlende bred vifte af farver & former - Heliconius erato f.eks findes der ikke mindre end 29 geografiske- eller klimaformer, der hver især svarer næsten præcis i farve og mønster til en "søster" underart af Heliconius melpomene der flyver i samme område.

Heliconius sara. Coroico 1900 m.h. d. 6 February 2012. Photographer Lars AndersenHeliconius sara. Coroico 1900 m.h. d. 6 February 2012. Photographer Lars Andersen

Sara Longwing, Heliconius sara. Coroico elevation 1900 m. 6 february 2012. Photographer; Lars Andersen

Variations

There are often striking differences between the different forms of each species, as can be seen in the illustrations here. In some subspecies of Heliconius erato the cream patch on the forewing is reduced to a group of dashes in the shape of a claw. In others it is replaced by a pair of large orange patches or may even be entirely absent. The basal area of the forewings is usually red, but may be unmarked in some races.

The hindwing markings may be red, orange or cream, either in the form of radiating lines, or as a solid median band.

Perhaps the most dramatic variety is H. cyrbia, which has a lurid pink band across the forewings, white submarginal rays on the hindwings, and a beautiful metallic blue sheen across the entire wing surface.

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Variationer

Der er ofte markante forskelle i mellem de forskellige former hos hver art. Hos Heliconius erato forskellige underarter er den beigefarvet plet i ydrefelt på forvinge overside reduceret til en gruppe streger i form af en klo. På andre underarter er det erstattes af et par store orangerøde pletter i ydre og inderfelt, eller måske mangler pletter helt. Rodfelt på forvinge er normalt rød eller med røde striber, men kan være helt mørk/sort.

På bagvinge overside som er sort, kan pletter eller bånd være rød, orange eller beigefarvet, enten i form af udstrålende linjer eller som et bånd igennem ydrefelt.

Måske den mest dramatiske form findes hos arten H. cyrbia, som har lyserøde bånd på tværs af forvinge, hvide submarginal stråler på bagvinge, samt et smuk metallisk blåt skær over hele overside.

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Heliconius erato. Coroico 1300 m.h. d. 9 february 2012. Photographer Lars AndersenHeliconius erato. Coroico 1300 m.h. d. 9 february 2012. Photographer Lars Andersen

Common Longwing, Heliconius erato venustus (Salvin, 1871) female with pollen on proboscis. Coroico, Yungas, elev. 1300 m. 9th February 2012. Photographer; Lars Andersen

Common Longwing, Heliconius erato

Common Longwing, Heliconius erato (Linnaeus, 1758) is probably the commonest and most widespread Heliconiini. It's 29 subspecies are distributed across the neotropical region from southern Texas, Mexico to Brazil, Bolivia, Paraguay and Missiones in northern Argentina.

The illustrated subspecies venustus (Salvin, 1871) is found in cloud forest in the Andes eastern mountain range of Yungas, northern Bolivia, elevation 1000 to 18oo m.

Subspecies luscombei (Lamas, 1976) is confined to the Amazonian lowlands of south-eastern Peru, while microclea (Kaye, 1907) is restricted to mid-elevation sites in central Peru, and petiverana (E. Doubleday, 1847) is Central American subspecies.

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Almindelig Langvinge, Heliconius erato

Almindelig Langvinge, Heliconius erato (Linnaeus, 1758) er nok den mest almindeligste og udbredte blandt Langvinger, Heliconiini.

Heliconius erato er delt op i 29 forskellige underarter, som er udbredt på tværs af neotropical regionen fra Mexico til Brasilien, Bolivia, Paraguay og Missiones i det nordlige Argentina.

Den viste underart venustus (Salvin, 1871) findes i tågeskovene i Yungas, Bolivia i 1000 til 1800 m.h.

Underarten luscombei (Lamas, 1976) findes i Amazonas lavland i det sydøstlige Peru, mens microclea (Kaye, 1907) findes i tågeskovene i 1000 til 2000 m.h. i det centrale Peru, og petiverana (E. Doubleday, 1847) findes udbredt i det sydøstlig Texas, det østlige Mexico til Panama.

Caranavi 7 February 2009. Photographer; Lars AndersenCaranavi 26 January 2009. Photographer; Lars Andersen

Common Longwing sitting on a dead Opossum. Caranavi, Yungas, Bolivia d. 7 February 2009. Photographer; Lars Andersen

Habitats

This species occurs commonly at elevations between 0-1800m on both sides of the Andes. It can be seen flying in two's or three's around clearings, and along roads and tracks through primary forest. It is commoner however in secondary forest, and can be found flying in coffee plantations, gardens, orchards and along roadsides and forest edges.

In regions where seasonality is pronounced, it is common in forest / pasture mosaics during the rainy season, but abandons these in favour of more heavily forested areas in the dry season.

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Levesteder

Denne art optræder almindeligvis i højder mellem 0-1800m på begge sider af Andesbjergene. Den optræder enkeltvis a 2 til 3 stk. omkring i lysninger, og langs veje og spor gennem primære skov. Den er mest almindelig i sekundær skov, og kan findes flyve i kaffeplantager, haver, frugtplantager og langs vejkanter og skovbryn.

I regioner hvor sæsonudsving er udtalt, er den almindelig i skov med græs mosaikker i regntiden, men opgiver disse til fordel for mere tæt skov i den tørre årstid.

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Heliconius erato. Coroico 1300 m.h. d. 19 january 2012. Photographer Lars AndersenHeliconius erato. Coroico 1300 m.h. d. 19 january 2012. Photographer Lars Andersen

Common Longwing, Heliconius erato mating on chrysalis. Coroico elevation 1300 m. 19 january 2012. Photographer; Lars Andersen

Lifecycle

The eggs are are round, slightly elongated with ribs and flat top, yellow, and placed individually on the leaf tips of Purple Passion, Passiflora menispermifolia and Orange-Petaled Passion, P. oerstedii.

When the caterpillars are fully grown they are white, dotted with black and have branched black spines on the back and sides. The head is straw coloured with a pair of recurved black spines. The caterpillars are aggressively cannibalistic.

The chrysalis is brown with golden spots on the abdomen and thorax. It has short black spikes on the abdomen and along the costa of the wing cases. The head is bifid, with the labial palpi extended and twisted. The overall impression is of a decaying dead twisted leaf, hanging from a stem.

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Livscyklus

Æggene er gule, og lægges enkeltvis på Lilla Passionsblomst, Passiflora menispermifolia & P. oerstedii blade.

Når larven er fuldt udvokset, er de hvide med sorte prikker og har forgrenet sorte pigge på overside og siden af de rudimentære segmenter (kropsled). Hovedet er stråfarvet med et par tilbagebøjede sorte pigge. Larver er aggressivt kannibalistisk.

Puppen er brun med gyldne pletter på maven og brystkassen. Den har korte sorte pigge på maven og langs krop og vingeskede. Hovedet har lange palpeskeder der er udvidet og snoet. Den ligner et rådnende snoet dødt blad, hængende fra en stilk.

Heliconius melpomene penelope. Coroico, Yungas, Bolivia d. 23 januar 2006. Fotograf: Lars Andersen

Postman, Heliconius melpomene penelope sitting on Psiguria flowers.
Coroico, Yungas, Bolivia d. 23 januar 2006. Photographer; Lars Andersen

Caranavi, Yungas, Bolivia d. 8 January 2010. Photographer: Lars Andersen

Wallace's Longwing, Heliconius wallacei. Caranavi, Yungas, Bolivia 8 January 2010. Photographer; Lars Andersen

Caranavi, Yungas, Bolivia d. 23 January 2010. Photographer: Lars AndersenCaranavi, Yungas, Bolivia d. 23 January 2010. Photographer: Lars Andersen

Tiger Mimic Longwing, Heliconius numata sitting on Red-Sage, Lantana camara flowers.
Caranavi, Yungas, Bolivia d. 23 January 2010. Photographer; Lars Andersen

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Agraulis vanillae laying eggs on passiflore. Coroico, Yungas, Bolivia. d. 23 january 2007. Photographer: Lars Andersen

Gulf Fritllary, Agraulis vanillae laying eggs on Passiflora. Coroico, Yungas, Bolivia d. 23 january 2007. Photographer; Lars Andersen

Adult behaviour

Heliconius butterflies are characterised by having a very delicate fluttering flight, particularly when hovering around flowers. They visit Hamelia, Lantana and Palicourea flowers for nectar.

Unlike other butterflies, Heliconius females feed on pollen as well as nectar. Studies of Heliconius ethilla have shown that females deprived of pollen can only produce about 15% of the number of eggs laid by females that have access to it. This probably applies equally to other Heliconius species including Postman, H. melpomene. The pollen from Psiguria, Anguria and Gurania flowers provides amino acids that can't be obtained from nectar or other sources, and contributes greatly to the longevity of the butterflies - some Heliconius species are known to live for up to 9 months as adults.

Studies have shown that Heliconius butterflies have home ranges within which they can memorise the locations of nectar and pollen sources, host plants and communal roosting sites. They are able to plan the most efficient route by which to visit all nectar / pollen sources in the vicinity by using simple calculations akin to what mathematicians call the "travelling salesman algorithm". Erlich & Gilbert demonstrated that individual butterflies memorise the location of particular Psiguria plants, which they visit daily, following a predefined circuit through the forest.

In the genus Heliconius most species rely entirely on airborne chemicals to locate mates. Males of Heliconius hecale, H. ismenius and H. cydno are attracted by pheromones to the pupae of conspecific females. The day before emergence a female pupa will usually have several males in close attendance. A frantic battle takes place the instant she hatches, as the males all struggle to copulate with her, not even allowing her time to expand and dry her wings. In some other Heliconius species such as H. hecalesia, H. hewitsoni, H. erato, H. charithonia and H. sara the males don't even wait until the female emerges. Instead they physically break open her pupa and copulate as soon as her genitalia are accessible.

Besides Heliconius males visits flowers, visiting also sandy banks along small streams, where they suck for minerals. They're also sucking on dead small mammals, as well as road kill.

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Heliconius telesiphe on Heliotropium flowers. Coroico 1300 m.h. d. 14 february 2012. Photographer Lars AndersenHeliconius telesiphe on Heliotropium flowers. Coroico 1300 m.h. d. 14 february 2012. Photographer Lars AndersenHeliconius telesiphe on Heliotropium flowers. Coroico 1300 m.h. d. 14 february 2012. Photographer Lars Andersen

Telesiphe Longwing, Heliconius telesiphe on Heliotropium flowers. Coroico 1300 m.h. d. 14 february 2012. Photographer; Lars Andersen

Adfærd

Heliconius sommerfugle er karakteriseret ved at have en meget følsom flagrende flugt, især når de flyver omkring blomster. De søger især nektar på Hamelia, Lantana og Palicourea blomster. I modsætning til andre sommerfugle, lever Heliconius hunner af pollen udover nektar. Undersøgelser har vist, at hunner frataget pollen kun kan producere ca. 15% af æg de normal lægger.

De udskiller pollen fra Psiguria, Anguria og Gurania blomster i regnskoven, pollen indsamlet fra blomsterne, behandles af hunnerne til at udvinde aminosyrer, som øger levetiden og sætte dem i stand til at producere æg i op til 9 måneder.

Undersøgelser har vist, at Heliconius sommerfugle kan huske ruter i tidsbegrænsede intervaller, inden for hvilke de kan huske de placeringer af nektar og pollen kilder, værtsplanter og fælles rastepladser. De er i stand til at planlægge den mest effektive rute, for at besøge alle nektar / pollenkilder i nærheden ved hjælp af enkle beregninger beslægtet med, hvad matematikere kalder "rejsende sælger algoritme".

Forskerne Erlich & Gilbert undersøgelser viser at de enkelte sommerfugle kan huske blomstrene Psiguria planter placering, som de besøger dagligt, efter en foruddefineret rute gennem skoven.

I slægten Heliconius vil de fleste arter udelukkende forlade sig på luftbårne kemikalier for at lokalisere artsfæller. Hos Heliconius hecale, H. ismenius og H. cydno hanner er tiltrukket af feromoner til uklækkede hun pupper. Dagen før puppen klækker, vil flere hanner søge puppe i tæt fremmøde. En hektisk kamp finder sted i det øjeblik, hun klækker, da alle hannerne kæmper for at parre sig med hende, de tillader hende ikke engang tid til at udvide og tørre sine vinger. Hos nogle arter som Heliconius hecalesia, H. hewitsoni, H. erato, H. charithonia og H. sara vil hannerne ikke engang vente til hun klækker. De vil i stedet fysisk åbne hendes puppe og parre, så snart hendes genitalier er tilgængelige.

Heliconius hanner besøger udover blomster, sandbanker langs små vandløb, hvor de suger mineraler. De sidder også og suger på døde små pattedyr, samt roadkill.

Caterpilar in Nymphalidae fam. Caranavi 605 m. Yungas, Bolivia d.  2 January 2014. Photographer:  Ole Andersen

Heliconius erato adults roost gregariously overnight, hanging in clusters of up to 10 from dry stems, usually quite close to the ground.

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Heliconius erato imago overnatter flere sammen op til 10 stk. hængende i små klynger på tørre stængler, normalt ganske tæt på jorden.

Her er der tale om forsvar! En omvandrene kaktus. Caranavi, Yungas, Bolivia d. 12 Februar 2006. Fotograf: Peter Møllmann

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Purple Passion, Passiflora menispermifolia.  Rio Zongo,  between Caranavi and Guarnay, Yungas. d. 26 january 2008. Photographer: Lars Andersen

Purple Passion, Passiflora menispermifolia. Rio Zongo, Caranavi, Yungas, Bolivia d. 26 january 2008. Photographer; Lars Andersen

Passiflora, known also as the passion flowers or passion vines, is a genus of about 500 species of flowering plants, the namesakes of the family Passifloraceae.

They are mostly vines, with some being shrubs, and a few species being herbaceous.

Passion fruit is a 4-6 centimeters round fruit. The shell is waxy and play in beautiful brown violet and green colors. The jelly-like flesh inside is filled with tiny black seeds.

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Passiflora, også kendt som Passionsblomst, er en slægt af omkring 500 arter af blomstrende planter, familien Passifloraceae.

De er for det meste kravlende slyngplanter, med nogle bliver buske, og nogle få arter er urteagtige.

Pasionsfrugt er en 4-6 centimer rund frugt. Skallen er voksagtig og spiller i kønne brunviolette og grønne farver. Det geleagtige frugtkød indeni er fyldt med små sorte frø.

Coroico. Yungas, Bolivia 20 january 2005. Photographer; Lars Andersen

Passionfruit or Maracuja, Passiflora edulis. Coroico. Yungas, Bolivia 20 january 2005. Photographer; Lars Andersen

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Mechanitis mazaeus holmgreni. Caranavi d. 26 January 2010. Photographer; Lars AndersenTiger mimic Longwing, Heliconius numata mirus (Weymer, 1894), en  Heliconidae som ligner Svalehalen til højre. Caranavi, Yungas, Bolivia d. 30 januar 2007. Fotograf: Lars AndersenBlack Mimic Swallowtail, Pterourus bachus chrysomelus (Rothschild & Jordan, 1906) , en Svalehale som deler udseende med den forgående heliconidae, på vejen imellem Yolosa og Caranavi, Yungas, Bolivia. 800 m.h. 11 Januar 2005. Fotograf: Lars Andersen

Mazaeus Tigerwing, Mechanitis mazaeus holmgreni (Bryk, 1953) Tiger Mimic Longwing, Heliconius numata mirus (Weymer, 1894) Black Mimic Swallowtail, Pterourus bachus chrysomelus (Rothschild & Jordan, 1906)
Tribe Ithomiini, subfamily: Danainae, family Nymphalidae. Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae. Tribe Papilionini, subfamily: Papilioninae, family Papilioidae.
Caranavi, Yungas, Bolivia d. 26 January 2010. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 30 January 2007. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 11 january 2005. Photographer; Lars Andersen
 

Mimicry

Heliconius butterflies have been a subject of many studies, due partly to their abundance and the relative ease of breeding them under laboratory conditions, but also because of the extensive mimicry that occurs in this group. From the nineteenth century to the present-day, their study has helped scientists to understand how new species are formed and why nature is so diverse. In particular, the genus is suitable for the study of both Batesian mimicry and Müllerian mimicry.

Mimicry, a generic imitation of one or more other species building features, coloration and sometimes behavior and fragrances.

The term is used most frequently on systems where the models are poisonous or otherwise dangerous: Nontoxic species some protection against predators who have learned to recognize and avoid the venomous species (Bates's mimicry, after the English naturalist and explorer Henry Walter Bates, 1825-1892) and toxic species reach among themselves to predators quickly learn to recognize them than if the toxic species were taught separately (Müller mimicry, after the German zoologist Fritz Müller, 1821-97).

Because of the type of plant material that Heliconius caterpillars favor and the resulting poisons they store in their tissues, the adult butterflies are usually unpalatable to predators.

This warning is announced, to the mutual benefit of both parties, by bright colors and contrasting wing patterns, a phenomenon known as aposematism. Heliconius butterflies are thus Müllerian mimics of one another, and are also involved in Müllerian mimicry with various species of Ithomiini, Danaini, Riodinidae (Ithomeis and Stalachtis) and Acraeini as well as Pericopine Arctiid moths. They are probably the models for various palatable Batesian mimics, including Black Mimic Swallowtail, Pterourus bachus and various Phyciodina.

One may wonder how animals that mimic other animals at all has occurred. The answer is evolution. Animals are changing from generation to generation, like all other living animals. These changes are usually so small that they are difficult to observe, but they can completely change the appearance and behavior of the animals for periods of over thousands or millions of years. This constant development called evolution.

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Cattleheart White, Archonias brassolis cutila (Fruhstorfer, 1907). Caranavi, Yungas, Bolivia. d. 30 januar 2006. Fotograf: Lars AndersenCaranavi, Yungas, Bolivia d. 12 Januar 2005. Fotograf: Lars AndersenEresia datis corybassa. Caranavi, Yungas. d. 9 february 2007. Photographer: Lars AndersenCaranavi, Yungas, Bolivia. d. 6 February 2007. Photographer: Lars Andersen

Cattleheart White, Archonias brassolis cutila (Fruhstorfer, 1907). Common Longwing, Heliconius erato venustus (Salvin, 1871) Hewitson's Mimic, Eresia datis corybassa (Hewitson, 1874) Mimic Tiger-Moths, Chetone phyleis (Druce, 1885)
Tribe Pierini, subfamily: Pierinae, family Pieridae. Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae. Tribe Melitaeini, subfamily Nymphalinae, family Nymphalidae. Tribe Pericopini, subfamily: Arctiinae, family Erebidae.
Caranavi, Yungas, Bolivia d. 30 january 2006. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 12 january 2005. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 9 february 2007. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 6 february 2007. Photographer; Lars Andersen
 

Mimicry

Ordet "mimicry" betyder, at noget kopieres så nøjagtigt så muligt. Når man taler om dyr, beskriver det en lighed mellem to dyr, som ikke er i nær familie. Der findes forskellige former for mimicry.

Mimicry, en arts efterligning af en eller flere andre arters bygningstræk, farvetegning, samt bevægelser (mimik), adfærd og duftstoffer. Betegnelsen benyttes hyppigst om systemer, hvor modellerne er giftige eller på anden måde farlige: Ugiftige arter opnår en vis beskyttelse mod rovdyr, der har lært at genkende og undgå de giftige arter (Bates' mimicry, efter Henry Walter Bates, 1825-1892 ), og giftige arter opnår indbyrdes, at rovdyr hurtigere lærer at genkende dem, end hvis de giftige arter skulle indlæres hver for sig (Müllers mimicry, efter den tyske zoolog Fritz Müller, 1821-97).

Heliconius sommerfugle har været genstand for mange undersøgelser, dels på grund deres store variationer i udseende og det er relativt nemt at opdrætte dem under laboratorieforhold, men også på grund af den omfattende mimicry, der opstår i denne gruppe. Fra det nittende århundrede til i dag, har undersøgelser i denne gruppe af sommerfugle hjulpet forskerne til at forstå, hvordan nye arter dannes og hvorfor naturen er så forskelligartet. Især slægten Heliconius er egnet til undersøgelse af både Batesian mimicry og Müllerske mimicry.

Stærke farver bruges ikke altid til at imponere med. Nogle dyr bruger stærke farver som et advarselssignal. Det gælder bl.a. stærke farver som rød, gul eller orange i kombination med sort i div. mønstre. Disse farver sender et signal til rovdyrene om, at "jeg er giftig og smager ikke godt".

Sommerfuglene får giften ved at larverne lever på planter som er giftige. Giften koncentreres i sommerfuglelarverne og gør, at den færdigudviklede sommerfugl også er giftig.

Cattleheart White, Archonias brassolis cutila (Fruhstorfer, 1907). Caranavi, Yungas, Bolivia. d. 30 januar 2006. Fotograf: Lars Andersen. Caranavi, Yungas, Bolivia d. 29 januar 2008. Fotograf: Lars AndersenHeliconius melpomene penelope. Caranavi, Yungas, Bolivia d. 29 januar 2008. Fotograf: Lars AndersenHeliconius erato. Coroico 1300 m.h. d. 9 february 2012. Photographer Lars Andersen

Cattleheart White, Archonias brassolis cutila (Fruhstorfer, 1907). Hewitson's Mimic, Eresia datis corybassa (Hewitson, 1874) Postman, Heliconius melpomene penelope (Staudinger, 1894). Common Longwing, Heliconius erato venustus (Salvin, 1871) .
Tribe Pierini, subfamily: Pierinae, family Pieridae. Tribe Melitaeini, subfamily Nymphalinae, family Nymphalidae. Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae. Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae.
Caranavi, Yungas, Bolivia d. 30 january 2006. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 1 february 2008. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 29 january 2008. Photographer; Lars Andersen Coroico, Yungas, elev. 1300 m. 9th February 2012. Photographer; Lars Andersen
 
Heliconius sommerfugle er således Müllerske mimisk efterligner af hinanden, og er også involveret i Müllerske efterligning med forskellige arter som ikke er nært beslægtede bruger samme farver - de har lavet en "forsvarsklub". Hvor diverse sommerfugle som Ithomiini, Danaini, Riodinidae (Ithomeis og Stalachtis) samt Acraeini & Pericopine arctiid møl.

Heliconius sommerfugle er også modeller for forskellige velsmagende Batesian minicry/efterlignere (mimikere), herunder Black Mimic Swallowtail, Pterourus bachus (C. Felder & R. Felder, 1865) og forskellige Phyciodina dagsommerfugle i tribe Melitaeini.

De forskellige sommerfugle arter signalerer, at de er giftige ved at have advarselsfarver som rød og gul kombineret med sort. Hvis disse sommerfugle arter så meget forskellige ud, ville det være svært for rovdyr at lære at holde sig fra netop disse sommerfugle. Ved at anvende samme farver og mønster opnår sommerfuglene en bedre gennemslagskraft med det advarselssignal, de sender. Grupper af disse sommerfugle arter sætter sig til hvile i træer om natten. Denne bunke af stærke farver skræmmer de fleste fjender væk. En jæger, som f.eks. en fugl, et firben eller en abe, skal bare æde én af disse sommerfugle for at vide, at de smager afskyeligt.

Man kan undre sig over, hvordan dyr der efterligner andre dyr overhovedet er opstået. Svaret er evolution. Dyr forandrer sig fra generation til generation, ligesom alle andre levende væsener. Disse forandringer er for det meste så små, at de er svære at observere, men de kan fuldstændig forandre dyrenes udseende og adfærd i perioder på over tusinder eller millioner af år. Denne konstante udvikling kaldes evolution.

Caranavi, Yungas, Bolivia d. 12 Januar 2005. Fotograf: Lars AndersenXanthocles Longwing, Heliconius xanthocles hippocrene (H. Holzinger & K. Brown, 1982). Caranavi, Yungas, Bolivia d. 17 february 2007. Photographer: Lars AndersenCaranavi, Yungas, Bolivia d. 30 january 2007. Fotograf: Lars AndersenCaranavi, Yungas, Bolivia d. 30 january 2007. Fotograf: Lars Andersen

Common Longwing, Heliconius erato venustus (Salvin, 1871) Xanthocles Longwing, Heliconius xanthocles hippocrene (H. Holzinger & K. Brown, 1982). Common Longwing, Heliconius erato venustus (Salvin, 1871) Common Longwing, Heliconius erato venustus (Salvin, 1871)
Tribe Helconiini, Tribe Helconiini, Tribe Helconiini, Tribe Helconiini,
subfamily: Heliconiinae, subfamily: Heliconiinae, subfamily: Heliconiinae, subfamily: Heliconiinae,
family Nymphalidae. family Nymphalidae. family Nymphalidae. family Nymphalidae.
Caranavi, Yungas, Bolivia d. 12 january 2005. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 17 february 2007. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 30 january 2007. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 31 january 2007. Photographer; Lars Andersen

Pterourus zagreus chrysomelus. Caranavi d. 8 january 2010. Photographer; Lars Andersen Tropical Milkweed Butterfly,  Lycorea halia pales (C. Felder & R. Felder, 1862) Tribe Danaini. Caranavi d. 3 February 2009. Photographer; Lars AndersenHypothyris ninonia cornelie (Guérin-Méneville, 1844), Caranavi, Yungas. february 2007. Photographer: Lars AndersenGolden Longwing, Heliconius hecale zeus. Caranavi, Yungas, Bolivia d. 14 februar 2007. Fotograf: Lars AndersenEunice Crescent, Eresia eunice (Hübner, 1807). Caranavi, Yungas. February 2008. Photographer: Lars Andersen

Tiger-Mimic Swallowtail, Pterourus zagreus chrysoxanthus (Fruhstorfer, 1915). Tropical Milkweed Butterfly, Lycorea halia pales (C. Felder & R. Felder, 1862). Common Glassy Tiger, Hypothyris euclea nina (Haensch, 1905). Harmonia Tiger, Tithorea harmonas (Cramer, 1777).   Eunice Crescent, Eresia eunice (Hübner, 1807).
Tribe Papilionini, Tribe Danaini, Tribe Ithomiini, Tribe Ithomiini, Tribe Melitaeini,
subfamily: Papilioninae, subfamily Danainae, subfamily Danainae, subfamily Danainae, subfamily Nymphalinae,
family Papilioidae. family Nymphalidae. family Nymphalidae. family Nymphalidae. family Nymphalidae.
Rio Coroico. Yungas, Bolivia d. 8 january 2010. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 3 february 2009. Photographer; Lars Andersen Caranavi, Yungas, Bolivia february 2007. Photographer; Lars Andersen Caranavi, Yungas, Bolivia d. 14 februar 2007. Photograper; Lars Andersen Caranavi, Yungas, Bolivia d. 15 february 2008. Photographer; Lars Andersen

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Tiger Leafwing, Consul fabius quadridentatus (Butler, 1874). Caranavi, Yungas. d. 1 february 2006. Photographer: Lars Andersen

Tiger Leafwing, Consul fabius quadridentatus (Butler, 1874). Caranavi, Yungas d. 1 february 2006. Photographer; Lars Andersen

Tiger Leafwing, Consul fabius som tilhører Charaxinae/Anaeini Underfamile. Vingeoversiden er lys orange og sort mønster, med to gule bånd på tværs af de vinkelformede forvingehjørne, bagvingerne har haler. Denne sommerfugl er en mimicry, som på oversiden efterligner Heliconiinae. Og undersiden med de kryptiske mønstre efterligner et dødt blad. Æggene lægges på blade af forskellige arter af Peber, Piperaceae (Anisillo, Piper tuberculatum, Mexican Pepper Leaf, Piper auritum, Cow-foot Leaf, Piper umbellatum, etc.). Er udbredt fra det sydlige Mexico, Panama, langs Andesbjergenes østskråninger til Yungas, Bolivia og Amazonas bassinet til Trinidad.

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Dryas iulia alcionea. Caranavi, elev. 800 m. d.  26 January 2012. Photographer: Lars Andersen

Julia, Dryas iulia alcionea (Cramer, 1779). Caranavi, elev. 800 m. d. 26 January 2012. Photographer; Lars Andersen

Henry Walter Bates, 1825-1892

Henry Walter Bates was an english naturalist and explorer who gave the first scientific account of mimicry in animals. He was most famous for his expedition to the rainforests of the Amazon with Alfred Russel Wallace, starting in 1848. Wallace returned in 1852, but lost his collection on the return voyage when his ship caught fire. When Bates arrived home in 1859 after a full eleven years, he had sent back over 14,712 species (mostly of insects) of which 8,000 were new to science. Bates wrote up his findings in his best-known work: "The Naturalist on the River Amazons". As describes similarities between toxic and harmless insects that are now described as Bates' mimicry. The phenomenon was recognized immediately as a strong support of Darwin's and Wallace's selection theories, which was then new.

Bates’s discussion of mimicry was unobtrusively buried in his classic article on the Longwing, Heliconius butterflies of the Amazon, which were frequently mimicked by counterfeits so perfect that even Bates was unable to distinguish them in flight. The Leptalides, Dismorphia butterflies, although quite different structurally from the Heliconius, are especially proficient mimics. Other examples abound. In various parts of the world beetles, spiders, flies, and grasshoppers mimic ants. While spiders may have a body configuration resembling that of ants, other mimics may use optical illusions to produce the appearance of a narrow antlike waist. Certain bees on the banks of the Amazon, as Bates observed, are also mimicked; indeed, many moths and longicorn beetles in the tropics mimic bees, wasps, and other hymenopterous insects.

Granted that mimics are adaptations to their environment, the important question was why such remarkably close analogies exist, Bates ruled out direct action of physical conditions because in limited districts where these conditions were the same, the most widely contrasting varieties may be found co-existing. Likewise, sports (mutations) did not explain mimicry. To Bates in was quite clear that natural selection had produced these phenomena, “the selecting agents being insectivorous animals which gradually destroy those sports or varieties which are not sufficiently like [the protected species] to deceive them.” The closer the resemblance of the mimic to the original, the greater will be its protection. Imperfect copies will be eliminated slowly, unless they have some supplementary protection of their own.

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Henry Walter Bates, 1825-1892

Henry Walter Bates var en engelsk naturforsker og opdagelsesrejsende, der lavede den første videnskabelige undersøgelse af mimicry hos forskellige dyrearter. Han var mest kendt for sin ekspedition til regnskovene i Amazonas sammen med Alfred Russel Wallace, der starter i 1848. Wallace tog tilbage i 1852, men mistede sin samling på returrejsen, da hans skib brød i brand. Da Bates kom hjem i 1859 efter en fuld elleve år, havde han sendt over 14,712 arter (hovedsagelig af insekter), hvoraf 8.000 var nye for videnskaben tilbage til England. Bates skrev sine resultater i hans bedst kendte værk: "The Naturalist on the River Amazons" som beskrev lighed mellem giftige og harmløse insekter, der nu betegnes som Bates' mimicry. Fænomenet erkendtes straks som en stærk støtte for Darwins og Wallaces selektionsteorier, som dengang var nye.

Bates diskussion af mimicry blev ubemærket begravet i hans klassiske artikel om Heliconius sommerfugle i Amazonas, som ofte blev efterlignes af forfalskninger så perfekt, at selv Bates var ude af stand til at skelne dem under flyvningen. F.eks. Dismorphia sommerfugle, som er helt forskellige strukturelt fra Heliconius, er særligt dygtige efterligner (mimikere). Andre eksempler findes i overflod. I forskellige dele af verden findes der biller, edderkopper, fluer, og græshopper der efterligner myrer. Mens edderkopper kan have et organ konfiguration ligner den hos myrer, kan andre efterlignere bruge optiske illusioner til at producere udseendet af en smal myrerlignende talje Visse bier på bredden af Amazonas, som Bates bemærkede, også efterlignede, ja, mange møl og langhorned biller i troperne efterligne bier, hvepse, og andre insekter som forskellige hvepse arter.

Bates konkluderet at det var helt klart, at naturlig udvælgelse havde produceret disse fænomener, "de insektædende dyr, som gradvist udrydder de dårlige efterlignere som ikke er tilstrækkeligt lignende de giftige arter, for at bedrage fjender. Jo tættere ligheden af mimik til den oprindelige, jo større er dens beskyttelse. Ufuldkomne kopier vil langsomt blive fjernet, med mindre de har nogle supplerende beskyttelse teknikker.

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Chetone species. Caranavi d. 25 January 2010. Photographer; Lars Andersen

Mimic Moths, Chetone species.
Tribe Pericopini, subfamily: Arctiinae, family Erebidae.
Caranavi, Yungas, Bolivia d. 25 january 2010. Photographer; Lars Andersen
 

Fritz Müller, 1821-97

Johann Friedrich Theodor Müller, better known as Fritz Müller, and also as Müller-Desterro, was a German biologist who emigrated to southern Brazil, where he lived in and near the German community of Blumenau, Santa Catarina. There he studied the natural history of the Atlantic forest south of São Paulo, and was an early advocate of Darwinism. He lived in Brazil for the rest of his life. Müllerian mimicry is named after him.

Müller's great discovery concerned the resemblance between two or more unpalatable species which are protected from predators capable of learning. The protection is often a noxious chemical, perhaps gained from the larva eating a particular plant; or it may be a sting or other defence. It is an advantage for such potential prey to advertise their status in a way clearly perceptible to their predators; this is called aposematic or warning coloration. The principle is of wide application, but in Müller's case the prey were butterflies, and the predators usually birds or reptiles.

The aposematic colours are most often some combination of red, yellow, black, white, whereas palatable animals are usually cryptic. The noxious animals may display by slow flying, and in general are prominently visible. Noxious animals usually have thick, leathery cuticles through which, at certain points, they extrude noxious fluids when pecked; they will often survive a 'trial'.

In Müllerian mimicry an advantage is gained when unpalatable species resemble each other, especially when the predator has a good memory for colour (as birds, for instance, do have). Thus one trial may work to dissuade a bird from several species of butterfly which all fly the same 'flag'. Brazilian butterflies provide some of the most extraordinary examples of mimicry, and Müller, Bates and Wallace all had lengthy experience of this. All three traveller-naturalists believed firmly that such systems of mimicry could only come about by means of natural selection, and all of them wrote about it.

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Fritz Müller, 1821-97

Johann Friedrich Theodor Müller, bedre kendt som Fritz Müller, og også som Müller-Desterro, var en tysk biolog, der emigrerede til det sydlige Brasilien, hvor han boede i og nær den tyske samfund af Blumenau, Santa Catarina. Der han studerede naturhistorie i skovene tæt på Atlanterhavet skoven syd for São Paulo, og var en tidlig fortaler for "Darwinismen". Han boede i Brasilien for resten af hans liv. Müllers mimicry er opkaldt efter ham.

Müllers store opdagelse vedrørte ligheden mellem to eller flere ubehagelige arter, der er beskyttet mod rovdyr der evner til at lære ved gentagelser. Beskyttelsen er ofte en kemisk giftig forsvar, som div. sommerfugle larver får igennem at spise en bestemt plante; eller det kan være et stik eller en anden forsvar. Det er en fordel for en sådan potentiel bytte at annoncere deres status på en måde der let at opfatte for deres rovdyr; dette kaldes aposematic eller advarsel farvning. Princippet er bredt anvendelige, men i Müller's undersøgelser var byttet sommerfugle, og rovdyr normalt fugle eller krybdyr.

De aposematic farver er oftest en kombination af rød, gul, sort, hvid, mens velsmagende dyr er normalt kryptisk farvet. De giftig dyr kan vise ved langsom flyvende, og de er generelt er fremtrædende synlige. Giftige dyr har normalt tykke, læderagtige krop og vinger, hvor de gennem på visse punkter, kan safte giftige væsker eller dufte, når fugle eller krybdyr hakker i dem; sommerfugle vil ofte overleve sådan et "forsøg".

I Müllerian mimicry er en fordel opnået, når ubehagelige arter ligner hinanden, især når rovdyr har en god hukommelse til farve (som fugle, for eksempel, har). Et forsøg kan således arbejde for at afholde en fugl fra flere arter af sommerfugle, som alle flyver med det samme farver og mønstre "flag". I Brazilien findes der sommerfugle som viser nogle af de mest usædvanlige eksempler på mimik, og Müller, Bates og Wallace har alle lang erfaring med dette. Alle tre rejsende-naturforskere troede fast på, at sådanne systemer af mimicry kun kunne ske ved hjælp af naturlig selektion, og alle drog lignende konklusioner om dette.

Alcoche, elev. 600 m. d.  30 January 2012. Photographer: Lars Andersen

Here tree different Heliconiini species; Bottom left on ground a Juno Longwing, Dione juno (Cramer, 1779), center, the big orange butterflies are Julia, Dryas julia alcionea (Cramer, 1779). One little orange in the middle is Juliette, Euides aliphera (Godart, 1819). From the road between Alcoche and Guarnay, Yungas, Bolivia d. 30 january 2012. Photographer; Lars Andersen.

Coroico Valley, Yungas, Sitio 70

Enero - Febrero 2012


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Telesiphe Longwing, Heliconius telesiphe (E. Doubleday, 1847)  on Gurania spinulosa flowers. The old railroad / Kori Wayku inca trail, Yungas, elev. 2000 m. 23 February 2009. Photographer: Lars Andersen Angle-winged Telesiphe, Podotricha telesiphe (Hewitson, 1867). Cloudy mountainforest; The old railroad between Coroico and Coripata,Yungas. d. 24  February 2008. Photographer: Prem Roy

Telesiphe Longwing, Heliconius telesiphe (E. Doubleday, 1847) Angle-winged Telesiphe, Podotricha telesiphe (Hewitson, 1867)
Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae. Tribe Helconiini, subfamily: Heliconiinae, family Nymphalidae.
On Gurania spinulosa flowers. The old railroad / Kori Wayku inca trail, elev. 2000 m. Yungas, Bolivia d. 23 February 2009. Photographer; Lars Andersen Cloudy mountainforest; The old railroad / Kori Wayku inca trail, elev. 2000 m. Yungas, Bolivia d. 24 February 2008. Photographer; Prem Roy

Coroico elevation 1800 m.h. Yungas, Bolivia d. 14 february 2012. Photographer; Lars Andersen

Coroico elevation 1800 m.h. Yungas, Bolivia d. 14 february 2012. Photographer; Lars Andersen

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Coroico, Yungas, Bolivia. d. 11 februar 2006. Fotograf: Lars AndersenCoroico, Yungas, Bolivia d. 19 januar 2005. Fotograf: Lars Andersen

Gulf Fritillary, Agraulis vanillae . Coroico, Yungas, Bolivia 11 february 2006 and 19 january 2005. Photographer; Lars Andersen

Gulf Fritillary or Passion Butterfly, Agraulis vanillae (Linnaeus, 1758) Description: is orange on the upper side, and frontwing with three small black spots with white core in discalcell, and black ribs on the marginal band. Its the rearwings are buff, with large silvery spots. Wingspand: medium to large butterfly, with a wingspan of 60 to 95 mm.

Habitats: is commonly seen in parks and gardens, as well as in open country.

Foodplants and caterpilar: The larva is a caterpillar which grows to approximately 4 cm in length. It is bright orange in color and covered in rows of black spines. The spines are soft to the touch and do not sting, but the larva is poisonous if eaten. The larva feeds exclusively on species of passionflower, such as Maypop, Passiflora incarnata, Yellow Passionflower, P. lutea, and Running Pop, P. foetida.

Distribution: Its range extends from Argentina north through Central America, Mexico, and the Antilles and Bahamas, Caribbean to the southern United States, as far north as the San Francisco Bay Area on the west coast. It is occasionally farther north. In Yungas, Bolivia flying the ssp. maculosa (Stichel, 1980).

Easy to hold, so it is often seen in butterfly farms. It is seen in Nexø, Bornholm, Denmark on Butterfly Bush, Buddleia davidii d. August 8, 2003 by Tom Kristensen Nygaard. It has probably escaped from the Bornholms Sommerfuglepark in the nearby area?

Gulf Fritillary, Agraulis vanillae. Nexø, Bornholm, Denmark d. 3 august 2003. Photographer; Tom Nygaard KristensenGulf Fritillary, Agraulis vanillae. Nexø, Bornholm, Denmark d. 3 august 2003. Photographer; Tom Nygaard Kristensen

Gulf Fritillary, Agraulis vanillae. Nexø, Bornholm, Denmark d. 3 august 2003. Photographer; Tom Nygaard Kristensen

Coroico, Yungas, Bolivia d. 20 januar 2006. Fotograf: Lars AndersenCoroico, Yungas, Bolivia d. 20 januar 2006. Fotograf: Lars Andersen

Gulf Fritillary or Passion ButterflyAgraulis vanillae maculosa (Stichel, 1908).
Hotel Esmeralda, Coroico, Yungas, Bolivia d. 20 january 2006. Photographer; Lars Andersen
 
PassionssommerfuglAgraulis vanillae (Linnaeus, 1758) er almindeligt set i parker og haver, såvel som i det åbne land i subtropisk til Tropisk klima. Dens udbredelse strækker sig fra det nordlig Argentina, langs Andesbjergene og Amazonas i Sydamerika til Mellemamerika, Mexico og Caribien til det sydlige USA, så langt nord som San Francisco Bay Area på vestkysten, og på østkysten op til New York.

Den er set længere nordpå i Canada på træk i Aweme, Manitoba (Brooks, 1942) og i Shilo, Manitoba, set af George Holland den 10. juni 1991.

Larven lever på Passionsblomst, Passiflora incarnata, Gul Passionsblomst, P. lutea og Vild Maracuja, P. foetida.

Caranavi d. 31 january 2008. Photographer; Lars AndersenAgraulis vanillae laying eggs on passiflore. Coroico, Yungas, Bolivia. d. 23 january 2007. Photographer: Lars Andersen

Gulf Fritillary or Passion ButterflyAgraulis vanillae maculosa (Stichel, 1908).
Caranavi, Yungas, Bolivia 31 january 2008, and Coroico 23 january 2007. Photographer; Lars Andersen

Coroico, Yungas, Bolivia d. 11 februar 2006. Fotograf: Lars AndersenCoroico 26 January 2009. Photographer; Lars Andersen

Juno LongwingDione juno (Cramer, 1779).
Coroico, Yungas, Bolivia 11 february 2006 and 26 january 2009. Photographer; Lars Andersen

Mariposas de Bolivia

Enero 2010

por Peter Møllmann y Lars Andersen


Mariposas de Bolivia 

Enero - Febrero 2009

por Peter Møllmann y Lars Andersen



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