Overview

Brief Summary

Introduction

Introduction:

The "blue morpho butterflies" represent one of the best known and most spectacular genera of Nymphalidae. They are mostly large - some with wingspans up to 20 cm, and many species have irridescent scales that reflect a metallic blue color. The blue color is due to refraction of light by fine striations on the surfaces of the wing scales - there is no blue pigment. The undersides of most species are brown with a variety of eye spots and smaller ocelli - features that reveal their affinity with other satyrines.

The classic living Morpho experience is to observe males patrolling along forest trails and stream beds appearing as a series of metallic blue flashes as the sun catches their wings. Collectors often equip themselves with a card covered with blue aluminum foil to attract the attention of males, who may swoop in to defend their territory against the incursion of a perceived interloper.

While the most familiar species are large and blue, there is quite a range of diversity in the genus (which has led to the description of numerous subgenera). Some Morpho species (M. epistrophus, M. polyphemus) are white, others are dark brownish gray or tawny orange (M. hecuba, M. telemachus). Some Andean species are smaller and quite delicate (M. lympharis).

Unlike most other satyrine taxa, some Morpho larvae feed on dicots. Penz and DeVries (2002) note that this larval host switch appears to have occurred once within the genus, the more basal species retaining the ancestral monocot diet.

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Introduction

The "blue morpho butterflies" represent one of the best known and most spectacular genera of Nymphalidae. They are mostly large - some with wingspans up to 20 cm, and many species have iridescent scales that reflect a metallic blue color. The blue color is due to refraction of light by fine striations on the surfaces of the wing scales - there is no blue pigment. The undersides of most species are brown with a variety of eye spots and smaller ocelli - features that reveal their affinity with other satyrines.

The classic living Morpho experience is to observe males patrolling along forest trails and stream beds appearing as a series of metallic blue flashes as the sun catches their wings. Collectors often equip themselves with a card covered with blue aluminum foil to attract the attention of males, who may swoop in to defend their territory against the incursion of a perceived interloper.

While the most familiar species are large and blue, there is quite a range of diversity in the genus (which has led to the description of numerous subgenera). Some Morpho species (M. epistrophus, M. polyphemus) are white, others are dark brownish gray or tawny orange (M. hecuba, M. telemachus). Some Andean species are smaller and quite delicate (M. lympharis).

Unlike most other satyrine taxa, some Morpho larvae feed on dicots. Penz and DeVries (2002) note that this larval host switch appears to have occurred once within the genus, the more basal species retaining the ancestral monocot diet.

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Comprehensive Description

Nomenclature

The genus Morpho has been subject of an enormous quantity of taxonomic revisionary work at the generic/subgeneric and specific/subspecific levels. The nomenclature employed here is based on the recent checklist of Lamas (2004), which adopts a relatively conservative view, including all the species within a single genus, and synonymizing many variations/geographical races under a relatively limited number of species. There are considerable differences in the nomenclature employed by Lamas vs. Penz & DeVries. Andy Brower is responsible for any errors of cross-referencing of the names on this tree.

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Evolution and Systematics

Evolution

Discussion of Phylogenetic Relationships

View Morpho Tree

The phylogenetic hypothesis is based upon Penz & DeVries (2002) morphological cladogram, with updates of nomenclature from Lamas (2004). Non-monotypic subgenera that appear as monophyletic taxa in the cladogram are labelled. However, as Penz and Devries noted, the polyphyly of many other subgenera suggests that until the systematics of the genus is better understood, perhaps these names should be avoided.

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Functional Adaptations

Functional adaptation

Wing scales diffract and scatter light: Morpho butterflies
 

Wings of Morpho butterflies create color by diffracting and scattering light.

   
  "Brilliant iridescent colouring in male butterflies enables long-range conspecific communication and it has long been accepted that microstructures, rather than pigments, are responsible for this coloration. Few studies, however, explicitly relate the intra-scale microstructures to overall butterfly visibility, both in terms of reflected and transmitted intensities and viewing angles.

"Using a focused-laser technique, we investigated the absolute reflectivity and transmissivity associated with the single-scale microstructures of two species of Morpho butterfly and the mechanisms behind their remarkable wide-angle visibility. Measurements indicate that certain Morpho microstructures reflect up to 75% of the incident blue light over an angle range of greater than 100 degree in one plane and 15 degree in the other.

"We show that incorporation of a second layer of more transparent scales, above a layer of highly iridescent scales, leads to very strong diffraction, and we suggest this effect acts to increase further the angle range over which incident light is reflected.

"Measurements using index-matching techniques yield the complex refractive index of the cuticle material comprising the single-scale microstructure to be n=(1.56 plus or minus 0.01) + (0.06 plus or minus 0.01)i. This figure is required for theoretical modelling of such microstructure systems." (Vukusic et al. 1999:1403)

  Learn more about this functional adaptation.
  • Vukusic, P. 1999. Quantified interference and diffraction in single Morpho butterfly scales. Proceedings of the Royal Society B: Biological Sciences. 266(1427): 1403-1403.
  • Yoshioka, S.; Kinoshita, S. 2004. Wavelength-selective and anisotropic light-diffusing scale on the wing of the Morpho butterfly. Proceedings: Biological Sciences. 271(1539): 581-587.
  • Prum, R. O.; Quinn, T.; Torres, R. H. 2006. Anatomically diverse butterfly scales all produce structural colours by coherent scattering. Journal of Experimental Biology. 209(4): 748-765.
  • Vukusic, P. 2006. Structural colour in Lepidoptera. Current Biology. 16(16): R621-R623.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
                                        
Specimen Records:728Public Records:316
Specimens with Sequences:625Public Species:59
Specimens with Barcodes:599Public BINs:50
Species:129         
Species With Barcodes:114         
          
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Barcode data

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Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Morpho

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Wikipedia

Morpho

This article is about the genus. For particular species, see Blue Morpho (disambiguation). For the electronic security company, see Morpho (Safran).

A morpho butterfly may be one of over 29 accepted species and 147 accepted subspecies of butterflies in the genus Morpho. They are Neotropical butterflies found mostly in South America as well as Mexico and Central America.[1] Morphos range in wingspan from the 7.5 cm (3 inch) M. rhodopteron to the imposing 20 cm (8 inch) Sunset Morpho, M. hecuba. The name Morpho, meaning changed or modified, is also an epithet of Aphrodite and Venus.

Species[edit]

This list is arranged alphabetically within species groups.[2]

Subgenus Iphimedeia

Subgenus Iphixibia

Subgenus Cytheritis

Subgenus Balachowskyna

Subgenus Cypritis

Subgenus Bessonia

Subgenus Crasseia

Subgenus Morpho

Ungrouped:

Taxonomy and nomenclature[edit]

A multitude of names attach to the genus Morpho, which has also been variously divided into subgenera. As well as names at species and subspecies level there are hundreds of form, variety, and aberration names. Lamas[3] included all such species within a single genus, and synonymizing many names in a limited number of species. C.M. Penz & P.J. DeVries[4] have provided a phylogenetic analysis with different nomenclature. Other authorities accept many more species than either.[5]

Etymology[edit]

The genus name Morpho comes from an Ancient Greek epithet "μορφώ", roughly "the shapely one", for Aphrodite, goddess of love and beauty.

Coloration[edit]

Many morpho butterflies are colored in metallic, shimmering shades of blues and greens. These colors are not a result of pigmentation but are an example of iridescence through structural coloration: the microscopic scales covering the Morpho's wings reflect incident light repeatedly at successive layers, leading to interference effects that depend on both wavelength and angle of incidence/observance.[6] Thus the colors produced vary with viewing angle, however they are actually surprisingly uniform, perhaps due to the tetrahedral (diamond-like) structural arrangement of the scales or diffraction from overlying cell layers. The wide angle blue reflection property can be explained by exploring the nanostructures in the scales of the Morpho butterfly wings.[7] These optical active structures integrate three design principles leading to the wide angle reflection: alternative lamellae layers, Christmas tree-like shape, and zigzag pattern of the ridges. The reflection spectrum is found to be broad (~ 90 nm) for alternating layers and can be controlled by varying the design pattern. The Christmas tree-like pattern helps to reduce the directionality of the reflectance by creating an impedance matching for blue wavelengths. In addition, the zigzag pattern of ridges destroys the unwanted interference for other wavelengths in wide angle. This structure may be likened to a photonic crystal. The lamellate structure of their wing scales has been studied as a model in the development of biomimetic fabrics, dye-free paints, and anti-counterfeit technology used in currency.

The iridescent lamellae are only present on the dorsal side of their wings, leaving the ventral side brown.

The ventral side is decorated with ocelli or eyespots. In some species, such as M. godarti, the dorsal lamellae are so thin as to allow the ventral ocelli to peek through. While not all Morphos have iridescent coloration, they all have ocelli. In most species only the males are colorful, supporting the theory that the coloration is used for intrasexual communication between males. The lamellae reflect up to 70% of light falling on them, including any ultraviolet (UV). The eyes of Morpho butterflies are thought to be highly sensitive to UV light and therefore the males are able to see each other from great distances. Some South American species are reportedly visible by the human eye up to one kilometre away.

There also exist a number of other species which are tawny orange or dark brown for instance (M. hecuba, M. telemachus). Some species are white principal among these being M. catenarius and M. laertes. An unusual species that is fundamentally white in coloration, but which exhibits a stunning pearlescent purple and teal iridescence when viewed at certain angles is the rare M. sulkowskyi. Some Andean species are small and delicate (M. lympharis). Among the metallic blue Morpho species, M. rhetenor stands out as the most iridescent of all, with M. cypris a close second. Indeed, M. cypris is notable in that specimens that are mounted in entomological collections will exhibit color differences across the wings if they are not 'set' perfectly flat.

Sexual dimorphism[edit]

The blue Morpho species exhibit sexual dimorphism. In some species (for instance M.adonis, M. eugenia, M. aega, M. cypris, and M. rhetenor) only the males are iridescent blue, the females are disruptively coloured brown and yellow. In other species (for instance M. anaxibia, M. godarti, M.didius, M. amathonte, and M. deidamia) the females are partially iridescent but less blue than the males.

Habitat[edit]

Atlantic Forest in Paraguay
  • Habitat: Primary forests of the Amazon and Atlantic. Also adapted to breed in a wide variety of other forested habitats, for instance the dry deciduous woodlands of Nicaragua and secondary forests.
  • Morphos are found at altitudes between sea level and about 1400 m.

Biology[edit]

  • Morphos are diurnal, as males spend the mornings patrolling along the courses of forest streams and rivers. They are territorial and will chase any rivals.Morphos typically live alone, excluding in the mating season.
  • The genus Morpho is palatable but some species (such as M. amathonte) are very strong fliers; birds—even species which are specialized for catching butterflies on the wing—find it very hard to catch them.[8] The conspicuous blue coloration shared by most Morpho species may be a case of Müllerian mimicry,[9] or may be 'pursuit aposematism'.[10]
  • The eyespots on the undersides of the wings of both males and females may be a form of automimicry in which a spot on the body of an animal resembles an eye of a different animal to deceive potential predator or prey species; to draw a predator's attention away from the most vulnerable body parts; or to appear as an inedible or even dangerous animal.[11]

Behavior[edit]

Morphos have a very distinctive slow, bouncy flight pattern due to the wing area being enormous relative to the body size.

Life cycle[edit]

The entire life cycle of the Morpho butterfly, from egg to death, is approximately 115 days.

caterpillars
pupae and emerging adult

The larvae hatch from pale green, dewdrop-like eggs. The caterpillars have reddish-brown bodies with bright lime-green or yellow patches on its back. Its hairs are irritating to human skin, and when disturbed it secretes a fluid that smells like rancid butter from eversible glands on the thorax. The strong odor is a defence against predators. They feed on a variety of plants. The caterpillar will molt five times before entering the pupal stage. The bulbous chrysalis is pale-green or jade-green and emits a repulsive, ultrasonic sound when touched.[12] It is suspended from a stem or leaf of the foodplant.

"The larvae live in nests on different forest – trees and especially on the climbing plants, but attack one another. There are about five moults. Larva cylindrical rather slender, somewhat thickened in the middle,tapering posteriorly. The last segment terminates in an indistinct tail-fork. The head is comparatively large and bears a pair of horizontal, conical processes, directed anteriorly, which are sometimes only rudimentary. Color always bright, sometimes variegated, yellowish with red-brown dorsal stripes or cross-shaped figures, back with long subdorsal tufts of bristles, of which the middle and posterior are in some cases gaily colored. Pupa similar to that of the genus Amathusia but more oval. Head with two tubercles, wing-cases distinctly projecting, abdomen sometimes belted with yellow ; color green or yellowish. Pupa on twigs or leaves attached by the abdomen but hanging free. The pupal stage lasts 20—30 days".[13]

The adults live for about two to three weeks. They feed on the fluids of fermenting fruit, decomposing animals, tree sap, fungi and nutrient rich mud.[14] They are poisonous to predators thanks to toxins they sequestered from plants they fed on as caterpillars.

The commoner Blue Morphos are reared en masse in commercial breeding programs. The iridescent wings are used in the manufacture of jewelry and as inlay in woodworking. Papered specimens are sold with the abdomen removed to prevent its oily contents from staining the wings. Significant quantities of live specimens are exported as pupae from several neotropical countries for exhibition in butterfly houses. Unfortunately, due to their irregular flight pattern and size, their wings are frequently damaged when in captivity.

Larvae host plants[edit]

Morpho larvae,variously according to species and region feed on Leguminosae, Gramineae, Canellaceae, Guttiferae, Erythroxylaceae, Myrtaceae, Moraceae, Lauraceae, Sapindaceae, Rhamnaceae, Euphorbiaceae, Musaceae, Palmae, Menispermaceae, Tiliaceae, Bignoniaceae, and Menispermaceae.

According to Penz and DeVries[4] the ancestral diet of larval Satyrinae is Poaceae or other monocots. Many Morphos have switched to dicots on several occasions during their evolutionary history but basal species have retained the monocot diet.

Collectors[edit]

Morpho butterflies, often very expensive, have always been prized by extremely wealthy collectors. Famous collections include those of the London jeweller Dru Drury and the Dutch merchant Pieter Teyler van der Hulst, the Paris diplomat Georges Rousseau-Decelle, the financier Walter Rothschild, the Romanov Grand Duke Nicholas Mikhailovich of Russia and the, English and German respectively, businessmen James John Joicey and Curt Eisner. In earlier years Morphos graced Cabinets of Curiosities "Kunstkamera" and Royal Cabinets of Natural History notably those of Tsar of Russia Peter the Great, the Austrian empress Maria Theresa and Ulrika Eleonora, Queen of Sweden. More famous is Maria Sibylla Merian who was not wealthy.

The people along the Rio Negro in Brazil once exploited the territorial habits of the Blue Morpho (M. menelaus) by luring them into clearings with bright blue decoys. The collected butterfly wings were used as embellishment for ceremonial masks. Adult Morpho butterflies feed on the juices of fermenting fruit with which they may also be lured. The butterflies wobble in flight and are easy to catch.

Photographs[edit]

Images of various varieties of morpho butterflies.

Illustrations of Morpho[edit]

See also[edit]

References[edit]

  1. ^ Le Moult (E.) & Réal (P.), 1962–1963. Les Morpho d'Amérique du Sud et Centrale, Editions du cabinet entomologique E. Le Moult, Paris.
  2. ^ Morpho, funet.fi
  3. ^ Lamas, G. (Ed.) 2004 Checklist: Part 4A. Hesperioidea-Papilionoidea. Gainesville, Florida: Association for Tropical Lepidoptera. ISBN 0-945417-28-4. [1]
  4. ^ a b Penz, Carla M.; DeVries, P. J. (2002). "Phylogenetic analysis of Morpho butterflies (Nymphalidae, Morphinae): Implications for classification and natural history". American Museum Novitates, (3374): 1–33. ISSN 0003-0082. 
  5. ^ https://sv.wikipedia.org/wiki/Morpho Species 2000 and Itis
  6. ^ P. Vukusic, J.R. Sambles, C.R. Lawrence, and R.J. Wootton (1999). "Quantified interference and diffraction in single Morpho butterfly scales". Proceedings of the Royal Society B 266 (1427): 1403–11. doi:10.1098/rspb.1999.0794. 
  7. ^ Siddique, R. H.; Diewald, S.; Leuthold, J.; Hölscher, H. (2013). Optics Express 21 (12): 14351–14361. doi:10.1364/OE.21.014351. 
  8. ^ Young, Allen M. (1971). "Wing colouration and reflectance in Morpho butterflies as related to reproductive behaviour and escape from avian predators". Oecologia 7 (3): 209–222. ISSN 0029-8549. 
  9. ^ Pinheiro, Carlos E. G. (1996). "Palatability and escaping ability in Neotropical butterflies: tests with wild kingbirds (Tyrannus melancholicus, Tyrannidae)". Biological Journal of the Linnean Society 59 (4): 351–363.  HTML abstract
  10. ^ Edmunds M. 1974. Defence in Animals: a survey of anti-predator defences. Harlow, Essex and NY: Longman. ISBN 0-582-44132-3. On pp. 255–256 there is a discussion of 'pursuit aposematism':

    "Young suggested that the brilliant blue colours and bobbing flight of Morpho butterflies may induce pursuit... Morpho amathonte is a very fast flier... It is possible that birds that have chased several unsuccessfully may learn not to pursue butterflies of that [type]... In one area, Young found that 80% of less brilliant species of Morpho had beak marks on their wings... but none out of 31 M. amathonte."

    "If brilliant colour was a factor in courtship, then the conflicting selection pressures of sexual selection and predator selection might lead to different results in quite closely related species."

  11. ^ Stevens, Martin (2005). "The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera". Biological Reviews 80 (4): 573–588. doi:10.1017/S1464793105006810. PMID 16221330. 
  12. ^ Greg Nussbaum Blue Morpho
  13. ^ Fruhstorfer, H. 1913. Family: Morphidae. In A. Seitz (editor), Macrolepidoptera of the world,vol. 5: 333–356. Stuttgart: Alfred Kernen.[2]
  14. ^ Blue Morpho Butterfly (Morpho peleides). Rainforest Alliance. Retrieved on 2011-10-17.

Further reading[edit]

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