Colias eurytheme

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Orange sulphur
McGeorge Colias eurytheme.JPG
Dorsal view
Orange sulphur in BBG (42933).jpg
Ventral view
Status TNC G5.svg
Secure  (NatureServe) [1]
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Pieridae
Genus: Colias
Species:
C. eurytheme
Binomial name
Colias eurytheme
Boisduval, 1852

Colias eurytheme, the orange sulphur, also known as the alfalfa butterfly and in its larval stage as the alfalfa caterpillar, is a butterfly of the family Pieridae, where it belongs to the lowland group of "clouded yellows and sulphurs" subfamily Coliadinae. It is found throughout North America from southern Canada to Mexico.

Contents

Other members of this lineage including the common or clouded sulphur ( C. philodice ) and C. eriphyle and C. vitabunda, which are often included in C. philodice as subspecies. Hybridization runs rampant between these, making phylogenetic analyses exclusively utilizing one type of data (especially mtDNA sequences) unreliable. Therefore, little more can be said about its relationships, except that it is perhaps closer to C. (p.) eriphyle than generally assumed, strengthening the view that the latter should be considered a valid species. [2]

The orange sulphur's caterpillars feed off various species in the pea family (Fabaceae) and are usually only found feeding at night. Occasionally this species multiplies to high numbers, and can become a serious pest to alfalfa (Medicago sativa) crops. The parasitoid wasp, Cotesia medicaginis can be used as a biocontrol agent against the caterpillars. [3]

Distribution

C. eurytheme butterflies can be found from southern Mexico to almost all throughout North America. Historically, they were distributed primarily in the western Nearctic, but were displaced to the east by logging and alfalfa field planting. [4]

Appearance

Wing pattern

Male C. eurytheme hindwings demonstrate an ultraviolet reflectance pattern while female C. eurytheme hindwings demonstrate ultraviolet absorbing patterns. According to studies, these ultraviolet reflecting wing scales found in males also contain pterin pigments that absorb wavelengths below 550 nm. Although this may seem paradoxical, the pterin pigments have been found to decrease the amount of diffuse ultraviolet reflectance that comes from the wing scales. By suppressing the diffuse ultraviolet reflectance, the directionality and spectral purity of the iridescence is heightened. In addition, the presence of the pterin pigments increases the signal's chromaticity and potential signal content, suggesting that these pigments are responsible for amplifying the contrast between ultraviolet reflectance and background colors as a male's wings move during flight. [5] Further studies have found that the ultraviolet reflectance signal is brightest within a wing beat cycle when viewed from directly above the male. This supports the idea that male wing color should be able to be readily distinguished from that of females and the visual background that consists mostly of UV-absorbing vegetation. [6]

Genetic inheritance

Studies have suggested that most of the genes controlling male courtship signals are inherited as a co-adapted gene complex on the X-chromosome. The X-chromosome carries most of the genes controlling production of 13-methyl heptacosane, the main component of pheromones involved in sexual selection, and the ultraviolet wing reflectance pattern. Expression of the ultraviolet wing reflectance pattern found in male C. eurytheme is controlled by a recessive allele on the X-chromosome. This trait is sex limited and not expressed in females of the same species. [7]

Reproduction

Reproductive behavior

Unlike that of many other butterfly species, the courtship of C. eurytheme is very brief and does not involve many elaborate displays. Mature female butterflies participate in mate selection by utilizing a specific refusal posture that prevents any undesired mating with both conspecific and non-conspecific males. [8]

These butterflies exhibit a polyandrous mating system. Upon mating, male C. eurytheme donate a nutritious spermatophore to the female, which will erode over time as nutrients are extracted for egg production and somatic maintenance. Females have a refractory period during which time they do not mate, but after they have depleted their spermatophore, they will search for another one and thus look for a new mate. In this mating system, females re-mate once every 4 to 6 days in summer, and mate a lifetime total of up to four times. [9]

Sexual selection

Male C. eurytheme have a visual cue (ultraviolet reflectance) and an olfactory cue (pheromones), both of which are suggested to be important in mate choice. Studies have suggested that pheromones may be more important in mediating female choice within a species, while ultraviolet reflectance may be more important in mediating female choice between species, such as between the very similar butterflies C. eurytheme and C. philodice. [10] The pheromone, located on the dorsal surface of the hindwing, consists of cuticular hydrocarbons n-heptacosane (C27), 13-methylheptacosane (13-MeC27), and possibly n-nonacosane (C29). [11] In addition, wing scales located on the dorsal wing surfaces in male C. eurytheme contain ridges with lamellae that produce iridescent ultraviolet reflectance via thin-film interference. [5]

C. eurytheme males rely on visual cues to locate and identify females. Instead of using chemical stimuli to find mates, males are attracted to the ultraviolet absorbing color of female hindwings. Studies have shown that males respond to paper dummies of the appropriate color and even attempt to mate with them. On the contrary, the ultraviolet reflection found on males strongly inhibits approaches from other males. This suggests that ultraviolet reflectance is also used by males as an inhibitory signal directed towards other males. [8]

Unlike sexual selection in males, visible color differences among males do not play an important role in mate selection by females. Females preferentially mate with males whose wings reflect ultraviolet light. [8] Studies have suggested that this trait was the strongest and most informative predictor of male courtship success. This may be because it has the potential to be an honest indicator of male condition, viability, and/or age. [10]

Due to the widespread cultivation of the alfalfa, the host plant for C. eurytheme and C. philodice, the species was able to expand their ranges across most of North America. These two species of sulphur butterflies have retained a large degree of genetic compatibility that allows them the produce viable and fertile offspring. [12] As a result of the recent sympatry and possible hybridization between these two species of sulphur butterflies, numerous studies have been conducted on intraspecific and interspecific mating. [13] In terms of mating under natural conditions, the males do not discriminate between the species, but females maintain nearly complete reproductive isolation. Studies suggest that the females do so by looking for the ultraviolet reflectance pattern on the dorsal wing surface of C. eurytheme males. [12] Therefore, it was suggested that C. eurytheme and C. philodice do not randomly mate with each other. Instead, mating was found to be positively assortive and mostly conspecific. [13]

Sexual selection theory

Previous studies have suggested that males make a nutrient investment during copulation. This idea agrees with the sexual selection theory, which predicts that females would act in ways to maximize the nutrient material they receive and predicts that males would act in ways to maximize the return on their investments. Studies support this theory by showing that younger males (males with less wing wear) are more successful in courtship than older males, males accepted by females are significantly less variable in size than males rejected by females, persistence increases a male's chance of copulating up to a point, and the size of females accepted by males is less variable than that of rejected females. [14] The amount of protein in a male's spermatophore is negatively correlated with age because it is more likely for older males to have mated previously. Females therefore prefer younger mates perhaps to secure large ejaculates, as smaller males and males that have mated previously produce smaller ejaculates. [9] The brightness of ultraviolet reflectance and pheromone descriptors, both important factors in mate selection, are also negatively correlated with age. However, variation between these two traits (visual and olfactory) is mostly uncorrelated. Since ultraviolet brightness emerges as the best predictor of male mating success, female preferences for brighter males may also indicate its relation to a material benefit. [10] In addition, studies have shown a longevity difference between virgin and mated females, suggesting a cost to mating. It is hypothesized that there is a toxic side effect of the male ejaculate. However, it is still unclear how this longevity cost influences the evolution of lifetime mating schedules. The supposed cost also does not affect the number of eggs a female lays in its lifetime. [15]

Related Research Articles

<span class="mw-page-title-main">Pheromone</span> Secreted or excreted chemical factor that triggers a social response in members of the same species

A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavior of the receiving individuals. There are alarm pheromones, food trail pheromones, sex pheromones, and many others that affect behavior or physiology. Pheromones are used by many organisms, from basic unicellular prokaryotes to complex multicellular eukaryotes. Their use among insects has been particularly well documented. In addition, some vertebrates, plants and ciliates communicate by using pheromones. The ecological functions and evolution of pheromones are a major topic of research in the field of chemical ecology.

<span class="mw-page-title-main">Green-veined white</span> Species of butterfly

The green-veined white is a butterfly of the family Pieridae.

<span class="mw-page-title-main">Pieridae</span> Butterfly family in superfamily Papilionoidea

The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia. Most pierid butterflies are white, yellow, or orange in coloration, often with black spots. The pigments that give the distinct coloring to these butterflies are derived from waste products in the body and are a characteristic of this family. The family was created by William John Swainson in 1820.

<span class="mw-page-title-main">Gulf fritillary</span> Sole species in brush-footed butterfly genus Agraulis

The Gulf fritillary or passion butterfly is a bright orange butterfly in the subfamily Heliconiinae of the family Nymphalidae. That subfamily was formerly set apart as a separate family, the Heliconiidae. The Heliconiinae are "longwing butterflies", which have long, narrow wings compared to other butterflies.

<i>Zerene eurydice</i> Species of butterfly

Zerene eurydice, the California dogface butterfly, belongs to the family Pieridae and is a sister genus to Colias.The Zerene eurydice and the Colias both share the "characteristic of having yellow-orange and black wing coordination." Additionally the,"Colias and Zerene eurydice males have bright UV patterns on their wings." There are only two species of the Zerene, the Zerene eurydice, and the Zerene cesonia, also known as the Southern dogface. A study that collected mitochondrial DNA from various Colias butterfly species found that Z. eurydice had decreased divergence from the ingroup, highlighting how closely related these two genera are. This species is endemic to California, and is California's state insect. The California dogface butterfly varies in its wing color and pattern.

<i>Anthocharis cardamines</i> Species of butterfly in the family Pieridae

Anthocharis cardamines, the orange tip, is a butterfly in the family Pieridae, which contains about 1,100 species. A. cardamines is mainly found throughout Europe and temperate Asia (Palearctic) The males feature wings with a signature orange pigmentation, which is the origin of A. cardamines' common name.

<i>Gonepteryx rhamni</i> Species of butterfly

Gonepteryx rhamni, commonly named the common brimstone, is a butterfly of the family Pieridae. It lives throughout the Palearctic zone and is commonly found across Europe, Asia, and North Africa. Across much of its range, it is the only species of its genus, and is therefore simply known locally as the brimstone. Its wing span size is 60–74 mm (2.4–2.9 in). It should not be confused with the brimstone moth Opisthograptis luteolata.

<i>Heliconius charithonia</i> Species of butterfly

Heliconius charithonia, the zebra longwing or zebra heliconian, is a species of butterfly belonging to the subfamily Heliconiinae of the family Nymphalidae. It was first described by Carl Linnaeus in his 1767 12th edition of Systema Naturae. The boldly striped black and white wing pattern is aposematic, warning off predators. It is the state butterfly of Florida.

<i>Colias croceus</i> Species of butterfly

Colias croceus, clouded yellow, is a small butterfly of the family Pieridae, the yellows and whites.

<i>Colias alfacariensis</i> Species of butterfly

Colias alfacariensis, Berger's clouded yellow, is a butterfly of the family Pieridae. It was separated from the pale clouded yellow, C. hyale, in 1905. Berger's clouded yellow is a Palearctic species (South and Central Europe, South Russia, Russian Far East, Siberia Central Asia and temperate China also Asia Minor, Caucasus and Transcaucasia.

<span class="mw-page-title-main">Queen (butterfly)</span> Species of butterfly

The queen butterfly is a North and South American butterfly in the family Nymphalidae with a wingspan of 80–85 mm. It is orange or brown with black wing borders and small white forewing spots on its dorsal wing surface, and reddish ventral wing surface fairly similar to the dorsal surface. The ventral hindwings have black veins and small white spots in a black border. The male has a black androconial scent patch on its dorsal hindwings. It can be found in meadows, fields, marshes, deserts, and at the edges of forests.

<i>Colias</i> Butterfly genus in family Pieridae

Colias is a genus of butterflies in the family Pieridae. They are often called clouded yellows in the Palearctic and sulphurs in North America. The closest living relative is the genus Zerene, which is sometimes included in Colias.

<i>Cosmophasis umbratica</i> Species of spider

Cosmophasis umbratica is a species of jumping spider found in South and Southeast Asia. They are members of the family Salticidae and the genus Cosmophasis. They are commonly spotted on green vegetation. C. umbratica shows extreme dimorphism when viewed under UV light: males reflect UV on all body parts that are displayed during intraspecific interaction, while females and juveniles do not reflect UV at all. It seems that C. umbratica uses this in sexual signaling. A similar phenomenon is found in some butterflies. For example, several species of Colias and Gonepteryx, both of the family Pieridae, also display sexual signaling.

<i>Pontia protodice</i> Species of butterfly

Pontia protodice, the checkered white or southern cabbage butterfly, is a common North American butterfly in the family Pieridae. Its green larva is a type of cabbage worm.

<i>Colias philodice</i> Species of butterfly

Colias philodice, the common sulphur or clouded sulphur, is a North American butterfly in the family Pieridae, subfamily Coliadinae.

<i>Colias tyche</i> Species of butterfly

Colias tyche, the Booth's sulphur or pale Arctic clouded yellow, is a butterfly in the family Pieridae. It is found from Baffin Island west along the Hudson Bay and arctic coasts of the Nunavut and Northwest Territories mainland and the southern tier of Arctic Islands to northern Yukon, Alaska, and Eurasia.

Butterflies, or members of the Papilionoidea superfamily, use two ultraviolet signals, UV reflectance or absorbance as a communication system. The ultraviolet region is the part of the electromagnetic spectrum between 10 nm and 400 nm in wavelength. Sensitivity to this region provides butterflies several benefits such as nectar guides for nectar, but it also provides a private communication channel unavailable to predators. With this secure line, butterflies are able to facilitate mating behavior and sex recognition.

<i>Colias alexandra</i> Species of butterfly

Colias alexandra, the Queen Alexandra's sulphur, Alexandra sulphur, or ultraviolet sulfur, is a butterfly in the family Pieridae found in western North America. Its range includes Alaska to the Northwest Territories and south to Arizona and New Mexico.

<span class="mw-page-title-main">Hair-pencil</span> Pheromone signaling structures in lepidopteran males

Hair-pencils and coremata are pheromone signaling structures present in lepidopteran males. Males use hair-pencils in courtship behaviors with females. The pheromones they excrete serve as both aphrodisiacs and tranquilizers to females as well as repellents to conspecific males. Hair-pencil glands are stored inside the male until courtship begins, at which point they are forced out of the body by sclerotized levers present on the abdomen. Coremata are very similar structures. Their exact definition is confused by early descriptions but they are more specifically defined as the internal, glandular, eversible structures that bear the hair-pencils and can be voluntarily inflated with hemolymph or air.

References

  1. ↑ "NatureServe Explorer 2.0 Colias eurytheme Orange Sulphur". explorer.natureserve.org. Retrieved 3 October 2020.
  2. ↑ Wheat, Christopher W. & Watt, Ward B. (2008). A mitochondrial-DNA-based phylogeny for some evolutionary-genetic model species of Colias butterflies (Lepidoptera, Pieridae). Mol. Phylogenet. Evol. 47(3):893-902. doi:10.1016/j.ympev.2008.03.013 (HTML abstract, supplement available to subscribers)
  3. ↑ Agriculture and Natural Resources, University of California. "UC Pest Management Guidelines: Alfalfa Caterpillar. UC ANR Publication 3430" . Retrieved 12 September 2017.
  4. ↑ Barton, Barb. "Colias eurytheme". Animal Diversity Web. University of Michigan Museum of Zoology. Retrieved 10 October 2013.
  5. 1 2 Rutowski, R.l, J. Macedonia, N. Morehouse, and L. Taylor-Taft. (2005). Pterin Pigments Amplify Iridescent Ultraviolet Signal in Males of the Orange Sulphur Butterfly, Colias Eurytheme. Proceedings of the Royal Society B: Biological Sciences 272(1578):2329-35.
  6. ↑ Rutowski, Ronald L., Joseph M. Macedonia, Justin W. Merry, Nathan I. Morehouse, Kasey Yturralde, Laura Taylor-Taft, Diann Gaalema, Darrell J. Kemp, and Randi S. Papke. (2007). Iridescent Ultraviolet Signal in the Orange Sulphur Butterfly (Colias eurytheme): Spatial, Temporal and Spectral Properties. Biological Journal of the Linnean Society 90(2):349-64.
  7. ↑ Grula, John W., and Orley R. Taylor. (1979). The Inheritance of Pheromone Production in the Sulphur Butterflies Colias eurytheme and C. Philodice. Heredity 42(3):359-71.
  8. 1 2 3 Silberglied, Robert E., and Orley R. Taylor. (1978). Ultraviolet Reflection and Its Behavioral Role in the Courtship of the Sulfur Butterflies Colias eurytheme and C. philodice (Lepidoptera, Pieridae). Behavioral Ecology and Sociobiology 3(3):203-43.
  9. 1 2 Kemp; Macedonia (2007). "Male mating bias and its potential reproductive consequence in the butterfly Colias eurytheme". Behavioral Ecology and Sociobiology. 61 (3): 415–422. doi:10.1007/s00265-006-0269-y. S2CID   4835212.
  10. 1 2 3 Papke, Randi S., Darell J. Kemp, and Ronald L. Rutowski. (2007). Multimodal Signalling: Structural Ultraviolet Reflectance Predicts Male Mating Success Better than Pheromones in the Butterfly Colias eurytheme L. (Pieridae). Animal Behavior 73:47-54.
  11. ↑ Sappington, T. W. (1990). Disruptive Sexual Selection in Colias Eurytheme Butterflies. Proceedings of the National Academy of Sciences 87(16):6132-5.
  12. 1 2 Grula, John W., and Orley R. Taylor. (1980). The Effect of X-Chromosome Inheritance on Mate-Selection Behavior in the Sulfur Butterflies, Colias eurytheme and C. Philodice. Evolution 34(4):688-95.
  13. 1 2 Taylor, Orley R., Jr. (1970). Random vs. Non-Random Mating in the Sulfur Butterflies, Colias eurytheme and Colias philodice (Lepidoptera: Pieridae). Evolution 26(3):344-56.
  14. ↑ Rutowski, Ronald L. (1985). Evidence for Mate Choice in a Sulphur Butterfly (Colias eurytheme). Zeitschrift für Tierpsychologie 70(2):103-14.
  15. ↑ Kemp, Darell J., and Ronald L. Rutowski. (2004). A Survival Cost to Mating in a Polyandrous Butterfly, Colias eurytheme. Oikos 105(1):65-70.
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