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== Description ==
== Description ==
The butterfly ranges over parts of [[North America|North]], [[Central America|Central]] and [[South America]], as well as the [[Caribbean]].<ref name="Savela Heliconius"/> In North America the butterfly is found in the southern parts of the [[United States]] including [[Florida]], [[Georgia (U.S. state)|Georgia]], [[Virginia]], [[North Carolina|North]] and [[South Carolina]].<ref name="LepIndex charithonia"/> In South and Central America, it has been recorded in [[Mexico]], [[Nicaragua]], [[Costa Rica]], [[Panama]], [[Colombia]], [[Ecuador]] and [[Venezuela]].<ref name="LepIndex charithonia"/><ref name="Savela Heliconius"/> Like Zebras they have a striped pattern giving them their name. Also like Zebras they each have a different stripe pattern.
The butterfly ranges over parts of [[North America|North]], [[Central America|Central]] and [[South America]], as well as the [[Caribbean]].<ref name="Savela Heliconius"/> In North America the butterfly is found in the southern parts of the [[United States]] including [[Florida]], [[Georgia (U.S. state)|Georgia]], [[Virginia]], [[North Carolina|North]] and [[South Carolina]].<ref name="LepIndex charithonia"/> In South and Central America, it has been recorded in [[Mexico]], [[Nicaragua]], [[Costa Rica]], [[Panama]], [[Colombia]], [[Ecuador]] and [[Venezuela]].<ref name="LepIndex charithonia"/><ref name="Savela Heliconius"/> Like Zebras they each have a unique striped pattern giving them their name.


== Life history ==
== Life history ==
The [[caterpillar]] feeds on Yellow Passionflower (''[[Passiflora lutea]]''), Corky-stemmed Passionflower (''[[Passiflora suberosa]]''), and Two-flower Passionflower (''[[Passiflora biflora]]''). The adults are unusual among butterflies in that they eat [[pollen]] as well as sip [[nectar]]. This ability contributes to their longevity—3 months as an adult.<ref>Scott, JA. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press.</ref> Because of their relatively long lifespan and their activity throughout the day, this is a popular species with butterfly houses. Another unusual feature is that adults roost in groups of up to 70, and return to the same roost each evening.
The [[caterpillar]] feeds on Yellow Passionflower (''[[Passiflora lutea]]''), Corky-stemmed Passionflower (''[[Passiflora suberosa]]''), and Two-flower Passionflower (''[[Passiflora biflora]]''). The adults are unusual among butterflies in that they eat [[pollen]] as well as sip [[nectar]]. This ability contributes to their longevity—3 months as an adult.<ref>Scott, JA. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press.</ref> Because of their relatively long lifespan and their activity throughout the day, this is a popular species with butterfly houses. Another unusual feature is that adults roost in groups of up to 70, and return to the same roost each evening.


== Behavior ==
==Defense Mechanism==
===Defense Against Predators===
Zebra Longwings use a variety of mechanisms in order to combat predation. With their [[aposematic]] coloration and unpalatability to predators, they exhibit [[Mullerian]] mimicry.<ref>{{cite journal|last=Kronforst|first=Marcus|coauthors=Theodore Fleming|title=Lack of genetic differentiation among widely spaced subpopulations of a butterfly with home range behaviour|journal=Heredity|year=2001|volume=86|pages=243-250}}</ref>
Zebra Longwings use a variety of mechanisms in order to combat predation. With their [[aposematic]] coloration and unpalatability to predators, they exhibit [[Mullerian]] mimicry.<ref>{{cite journal|last=Kronforst|first=Marcus|coauthors=Theodore Fleming|title=Lack of genetic differentiation among widely spaced subpopulations of a butterfly with home range behaviour|journal=Heredity|year=2001|volume=86|pages=243-250}}</ref>



Revision as of 21:46, 9 October 2013

Zebra Longwing
Dorsal view - feeding from a sage flower
Ventral view
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
H. charithonia
Binomial name
Heliconius charithonia
Synonyms
  • Heliconius charithonius

The Zebra Longwing or Zebra Heliconian (Heliconius charithonia)[1][2] is a species of butterfly belonging to the subfamily Heliconiinae of the Nymphalidae. It was declared the official butterfly for the state of Florida in the United States in 1996.

Description

The butterfly ranges over parts of North, Central and South America, as well as the Caribbean.[2] In North America the butterfly is found in the southern parts of the United States including Florida, Georgia, Virginia, North and South Carolina.[1] In South and Central America, it has been recorded in Mexico, Nicaragua, Costa Rica, Panama, Colombia, Ecuador and Venezuela.[1][2] Like Zebras they each have a unique striped pattern giving them their name.

Life history

The caterpillar feeds on Yellow Passionflower (Passiflora lutea), Corky-stemmed Passionflower (Passiflora suberosa), and Two-flower Passionflower (Passiflora biflora). The adults are unusual among butterflies in that they eat pollen as well as sip nectar. This ability contributes to their longevity—3 months as an adult.[3] Because of their relatively long lifespan and their activity throughout the day, this is a popular species with butterfly houses. Another unusual feature is that adults roost in groups of up to 70, and return to the same roost each evening.

Defense Mechanism

Zebra Longwings use a variety of mechanisms in order to combat predation. With their aposematic coloration and unpalatability to predators, they exhibit Mullerian mimicry.[4]

Mating System

Mating cues

Male butterflies exhibit a preference for visual, olfactory, tactile, or auditory cues during mating so that females are made more obvious.[5] In Heliconius charithonia, certain host plants provide these cues to males, thereby influencing the time and location of reproduction. This happens because as larvae damage the plant upon eating it, green-leaf volatiles, six carbon alcohols, aldehydes, and acetates, are released. They give olfactory cues to the male, thereby indicating the location of the pupae (mate). Since these pupae have camouflaged coloring and lack strong sexual pheromones, the olfactory cue given off from the damaged plant is necessary in order for the male to find a mate. Furthermore, the odors given off from the plant not only signal the location of pupae but also provide a trigger that induces learning the location of the plant for future copulations. This is possible due to the fact that the butterflies have a sophisticated spatial memory, as exhibited by their tendency to have specific nocturnal roosting locations and regular visitations of sites with abundant adult resources.[6]

A common problem amongst all butterflies is the ability of mates to discriminate between conspecifics and other butterfly species.[7] Although mistakes do occur, they are quite uncommon and males are able to distinguish, more often than not, between the emissions produced upon the larvae eating and other herbivores eating the plant. Due to this observation, it has been found that the zebra longwing also uses larval coloration and odors in order to find mates. Chemical analysis has demonstrated that the larvae release volatiles that are similar chemically to those emitted by the plant.[8]

Heliconius chartihonia also demonstrates pleiotropic mating cues, particularly in regards to mimicry. Pleiotropic mating cues have been simultaneously ecologically and sexually selected. Mimetic patterns have been used in cues to mate-finding as well as in predatory protection, leading to the coevolution of mate choice and assortative mating, a nonrandom mating pattern in which individuals choose mates with similar genotypes and/or phenotypes to themselves.[9] Polymorphism exists in mimetic patterns due to the fact that each morph mimics a different model.[10]

Pupal mating

Males perform precopulatory mate guarding behavior, in which males find and perch on pupae, followed by copulation with the female.[11] Upon reaching the pupae, males often have to compete in order to copulate with the female, who is teneral (freshly emerged). Typically, a male visits the same pupa for at least a week, during which time he periodically swarms it, fighting with other males over positioning. Fights consist of males fending off other males that attempt to land on the same pupa by opening their wings. If this does not work, the male tries to throw the intruder off with the pressure of his head and antennae. If more males attempt to swarm the pupa, the two original males will work together to attempt to fend off the others by simultaneously opening their wings, momentarily forgetting that they were originally competitors. Fights usually last one or two hours, but continue throughout the pupa’s development.

The act of pupal mating consists of the male inserting his abdomen into the pupa. If there is a second male, he will fend off other males by opening his wings during the copulation of the first male, rather than attempting to mate with the female himself by inserting his abdomen. After two or three hours of mating, the female comes out during which copulation stills occurs for another hour. During the process, females remain relatively still, with the exception of the spreading of their wings and the discharge of meconium. As the copulation proceeds, less and less males attempt to approach the female. However, if this does occur, the copulating male continues to fend them off by opening his wings. After copulation is done, the male and female sit side-by-side for some time. During this brief period, no other males attempt to mate with the female. Pupal mating arose once during the evolution of Heliconius, and these species form a clade on the evolutionary tree. Although pupal mating is observed quite frequently in insectaries, it is rarely seen in nature.[12]

Nuptial gifts in the form of the spermatophore

Males transfer a protein-rich spermatophore to females upon mating. Spermatophores are thought to be nuptial gifts, which serve a number of different functions. One of these is to provide chemicals (cyanogens) that protect the mother and future offspring from predators so that female and egg survival are enhanced. For females, this is beneficial because egg-laying causes a depletion of protein and her own defensive chemicals. Among nine Heliconius species studied, Heliconius chartihonia had the highest average cyanide concentration in its spermatophores.[13]

In most species of butterflies, pheromones play a role in courtship and mate recognition.[14] However, they can also play a role in deterring mates. Spermatophores are also thought to contain antiaphrodisiacs, which are pheromones that reduce the attractiveness of the females to subsequent males, thereby indicating that antiaphrodisiac evolution is driven by intrasexual selection. Because of this, they help reduce the occurrence of male harassment by already mated females. Furthermore, nonfertile sperm (apyrene) is added to spermatophores in order to increase the time of refractory periods. Overall, the transfer of antiaphrodisiacs is a mechanism for degrading female mating choice.[15]

It has been observed that complete spermatophore degradation to an orange or yellow substance occurs in a 2-week period. Pupal-mating butterflies like Heliconius charatonia are thought to be monandrous, but spermatophore degradation could undermine this idea since spermatophores are often examined to count the number of matings a female has performed. However, upon further observation, it was found that females rarely participate in more than one mating per lifetime.[16]

Sex ratio and distribution

Reproduction occurs year-round. Although at eclosion the ratio is highly female-biased, the sex ratio is overall male-biased (68% males). This is due to the fact that males typically stay near their natal sites in order to find a mate while females move around in order to find oviposition or feeding sites at various Passiflora plants. Because females are very mobile, males do not mate with relatives very often, and therefore inbreeding rates are very low.[17]

Taxonomic note

In some publications the butterfly is referred to as Heliconius charitonius, but this is either a lapsus calami (slip of the pen) or unjustified emmendation of the original name given by Linnaeus in 1767, Papilio charithonia. Cramer in 1777 was apparently the first to publish this incorrect name. Godman & Salvin in 1901 referred to the species as H. charithonie [sic].

See also

Gallery

References

  1. ^ a b c Card for charithonia in LepIndex. Accessed 3 August 2007.
  2. ^ a b c Marrku Savela's Website on Lepidoptera Heliconius, funet.fi
  3. ^ Scott, JA. 1986. The Butterflies of North America: A Natural History and Field Guide. Stanford University Press.
  4. ^ Kronforst, Marcus (2001). "Lack of genetic differentiation among widely spaced subpopulations of a butterfly with home range behaviour". Heredity. 86: 243–250. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ Douglas, Matthew M. The Lives of Butterflies. Ann Arbor: University of Michigan, 1986. Print.
  6. ^ Estrada, Catalina, and Lawrence E. Gilbert. "Host Plants and Immatures as Mate- searching Cues in Heliconius Butterflies." Animal Behaviour 80 (2010): 231-39. Web.
  7. ^ Boggs, Carol L., Ward B. Watt, and Paul R. Ehrlich. Butterflies: Ecology and Evolution Taking Flight. Chicago: University of Chicago, 2003. Print.
  8. ^ Estrada, Catalina, and Lawrence E. Gilbert. "Host Plants and Immatures as Mate- searching Cues in Heliconius Butterflies." Animal Behaviour 80 (2010): 231-39. Web.
  9. ^ Jiggins, Chris D., Igor Emelianov, and James Mallet. "Assortative Mating and Speciation as Pleiotropic Effects of Ecological Adaptation: Examples in Moths and Butterflies." Insect Evolutionary Ecology. By Mark Fellowes, G. J. . Holloway, and J. Roff. Wallingford, Oxfordshire: CABI Pub., 2005. 451-73. Print.
  10. ^ Scoble, M. J. The Lepidoptera: Form, Function and Diversity. [London]: Natural History Museum, 1995. Print.
  11. ^ Estrada, Catalina, and Lawrence E. Gilbert. "Host Plants and Immatures as Mate- searching Cues in Heliconius Butterflies." Animal Behaviour 80 (2010): 231-39. Web.
  12. ^ Sourakov, Andrei. "Pupal Mating in Zebra Longwing (Heliconius Charithonia): Photographic Evidence." News of the Lepidopterists' Society 50.1 (2008): 26-32. Print.
  13. ^ Cardoso, Márcio Zikán, and Lawrence E. Gilbert. "A Male Gift to Its Partner? Cyanogenic Glycosides in the Spermatophore of Longwing Butterflies (Heliconius)." Naturwissenschaften 94.1 (2006): 39-42. Print.
  14. ^ Douglas, Matthew M. The Lives of Butterflies. Ann Arbor: University of Michigan, 1986. Print.
  15. ^ Estrada, Catalina, Stefan Schulz, Selma Yildizhan, and Lawrence E. Gilbert. "Sexual Selection Drives The Evolution Of Antiaphrodisiac Pheromones In Butterflies." Evolution 65.10 (2011): 2843-854. Print.
  16. ^ Walters, James R., Christine Stafford, Thomas J. Hardcastle, and Chris D. Jiggins. "Evaluating Female Remating Rates in Light of Spermatophore Degradation in Heliconius Butterflies: Pupal-mating Monandry versus Adult-mating Polyandry." Ecological Entomology 37 (2012): 257-68. Print.
  17. ^ Fleming, Theodore H., David Serrano, and Jafet Nassar. "Dynamics Of A Subtropical Population Of The Zebra Longwing Butterfly Heliconius Charithonia (Nymphalidae)." Florida Entomologist 88.2 (2005): 169-79. Print.

External links