Overview

Brief Summary

North American Ecology (US and Canada)

Resident throughout the US and southern Canada, migratory to the north (Scott 1986). Habitats are OPEN AREAS at all altitudes. Host plants are usually herbaceous including many species, but mostly in one family, Leguminosae. Eggs are laid on the host plant singly. Individuals overwinter as 3rd or 4th instar larvae. There are multiple flights each year with the approximate flight time FEB1-NOV1 depending on latitude (Scott 1986).
  • Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
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Comprehensive Description

General Description

The combination of an orange upperside, halo-like ring around the discal spot of the hindwing underside, and row of submarginal spots will distinguish this species in most cases. Albino females are very similar to those of C. philodice, although slightly larger. There are no recognized subspecies.
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Distribution

Central Mexico north to central Canada (Opler 1999). It is uncertain how far north this species is able to overwinter.
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occurs (regularly, as a native taxon) in multiple nations

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National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Most of North America; rarer in subtropical Florida, Canada, Alaska, and the coastal Northwest. Nearly lacks any diapause but is very cold tolerant in all stages except egg, and this species occurs year round where the January mean is above zero Celsius or just below, e.g most of Kentucky, extreme southern Pennsylvania and perhaps half of New jersey in the east.. An abundant summer resident hundreds of km farther north.

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Geographic Range

Orange sulphurs historically were a western species in the Nearctic region, but moved eastward across North America during the late 1800’s due to logging and the planting of alfalfa fields. They now are found throughout North America to southern Mexico.

Biogeographic Regions: nearctic

  • Iftner, D., J. Shuey, J. Calhoun. 1992. Butterflies and Skippers of Ohio. Columbus, OH: Ohio Biological Survey Bulletin, Vol 9 No. 1.
  • Scott, J. 1986. The Butterflies of North America. Stanford, CA: Stanford University Press.
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Physical Description

Morphology

Physical Description

The upper surface of the wings is primarily orange, although some females are white. The underside of the hindwing has a silver spot encircled by two red rings and a satellite spot. The upper surface of the males’ wings reflect ultraviolet, which is caused by a recessive gene on the X chromosome. Orange sulphurs are strongly polymorphic, and the general practice is if a sulphur has any orange on the wings at all it is called an orange sulphur. The average wing length of males is 2.4 cm, with a range of 2.1 - –2.8 cm. Average females wing length is 2.6 cm, with a range of 2.3 - –3.1 cm.

The cream colored eggs are spindle shaped and turn crimson with age.

The larvae are green with a white lateral band and faint green dorsal lines.

Range wingspan: 4.1 to 6.2 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry ; polymorphic

Sexual Dimorphism: female larger; sexes colored or patterned differently

  • Opler, P., G. Krizek. 1984. Butterflies East of the Great Plains. Baltimore: John Hopkins University Press.
  • Shapiro, A. 1966. Butterflies of the Delaware Valley. American Entomological Society Special Publication.
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Type Information

Lectotype for Colias eurytheme Boisduval, 1852
Catalog Number: USNM
Collection: Smithsonian Institution, National Museum of Natural History, Department of Entomology
Sex/Stage: Male; Adult
Preparation: Pinned
Locality: California, United States
  • Lectotype: Annales de la Société Entomologique de France. 10 (2): 286.
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Ecology

Habitat

Found in open areas throughout the province, particularly roadsides and agricultural areas.
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Comments: Almost any kind of opening with legumes.

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This species can be found in most any open area, including vacant lots, pastures, open fields, roadsides, and clover and alfalfa fields.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: savanna or grassland

Other Habitat Features: urban ; suburban ; agricultural ; riparian

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Migration

Non-Migrant: No. All populations of this species make significant seasonal migrations.

Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

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Trophic Strategy

Larvae feed on a wide variety of legumes, particularly non-natives such as clover (Trifolium) and alfalfa (Medicago sativa) (Guppy & Shepard 2001). There are no larval records of this species for Alberta, but these plants are the most likely hosts. Larvae sometimes reach pest levels in the southern portions of the range (Layberry et al. 1998). Adults take nectar at legume flowers, including alfalfa, and males mud-puddle (Nielsen 1999).
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Food Habits

The hostplants are in the family Fabiacae. The main food plant of the caterpillars is alfalfa (Medicago sativa).

Adults will sip from mud puddles and take nectar from a variety of plant species, including alfalfa, clovers (Trifolium), milkweeds (Asclepias), selfheal (Prunella vulgaris), teasel (Depsacus sylvestris), peppermint (Mentha piperita), horseweed (Erigeron canadensis), purple coneflower (Echinacae pupurea), sunflower (Helianthus hirsutus), asters (Aster), and goldenrods (Solidago).

Plant Foods: leaves; nectar

Primary Diet: herbivore (Folivore , Nectarivore )

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Associations

Flowering Plants Visited by Colias eurytheme in Illinois

Colias eurytheme Boisduval: Pieridae, Lepidoptera
(observations are from Hilty, Clinebell, Fothergill & Vaughn; this butterfly is the Orange Sulfur; some observations by Robertson and Graenicher apply to the Orange Sulfur under Colias philodice as they did not distinguish between these two species)

Asteraceae: Aster novae-angliae sn (H), Aster subulatus sn (FV), Cirsium discolor sn (H), Echinacea pallida sn (Cl), Eupatorium altissimum sn (H), Liatris aspera sn (H), Oligoneuron rigidum sn (H), Silphium integrifolium sn (H), Taraxacum officinale sn (FV); Lamiaceae: Lamium amplexicaule sn (FV), Monarda fistulosa sn (Cl)

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Ecosystem Roles

Orange sulphurs serve as minor pollinators and prey for many species of predators.

Ecosystem Impact: pollinates

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Predation

Predators of all life stages of butterflies include a variety of insect parasatoids. These wasps or flies will consume the body fluids first, and then eat the internal organs, ultimately killing the butterfly. Those wasps that lay eggs inside the host body include species in many different groups: Ichneumonidae, Braconidae, Pteromalidae, Chalcidoidea, Encyrtidae, Eulophidae, Scelionidae, Trichogrammatidae, and others. Trichogrammatids live inside the eggs, and are smaller than a pinhead. Certain flies (Tachinidae, some Sarcophagidae, etc.) produce large eggs and glue them onto the outside of the host larva, where the hatching fly larvae then burrow into the butterfly larvae. Other flies will lays many small eggs directly on the larval hostplants, and these are ingested by the caterpillars as they feed.

Most predators of butterflies are other insects. Praying mantis, lacewings, ladybird beetles, assasin bugs, carabid beetles, spiders, ants, and wasps (Vespidae, Pompilidae, and others) prey upon the larvae. Adult butterflies are eaten by robber flies, ambush bugs, spiders, dragonflies, ants, wasps (Vespidae and Sphecidae), and tiger beetles. The sundew plant is known to catch some butterflies.

There are also many vertebrate predators including lizards, frogs, toads, birds, mice, and other rodents.

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Known predators

Colias eurytheme is prey of:
Orius insidiosus

Based on studies in:
USA: Illinois (Agricultural)

This list may not be complete but is based on published studies.
  • M. A. Mayse and P. W. Price, 1978. Seasonal development of soybean arthropod communities in east central Illinois. Agro-Ecosys. 4:387-405, from p. 402.
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Known prey organisms

Colias eurytheme preys on:
Glycine max

Based on studies in:
USA: Illinois (Agricultural)

This list may not be complete but is based on published studies.
  • M. A. Mayse and P. W. Price, 1978. Seasonal development of soybean arthropod communities in east central Illinois. Agro-Ecosys. 4:387-405, from p. 402.
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: > 300

Comments: Probably well over a million each summer, less than a million in mid winter.

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Global Abundance

>1,000,000 individuals

Comments: Could easily exceed 100,000,000,000 by September probably about 10% of that by February.

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Life History and Behavior

Behavior

Communication and Perception

Orange sulphurs find potential mates using their vision.

Communication Channels: visual ; tactile

Perception Channels: visual ; ultraviolet; tactile

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Adults feed mainly from nectar and mud. Males patrol for females (Scott, 1986).
  • Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
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Cyclicity

Two broods, flying in June and early August to early September.
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Life Cycle

The egg is initially white, turning red several days after being laid. Mature larvae are dark, velvety green with a red-bordered, white lateral line with yellow to red dashes (Guppy & Shepard 2001). Pupae are light green with a yellow lateral line and brown markings. This species fluctuates greatly in abundance from year to year, being rare or absent in some years and common in others; this is apparently the result of winter survival in areas to the south of Alberta, since the Orange Sulphur cannot survive the Alberta winters. Migrants of the Orange Sulphur appear in central Alberta in late June to early July; these migrants are larger than the summer brood they produce here, and are usually flight-worn by the time they reach Alberta. It is not known if individuals of the second brood attempt a southward migration, or if they perish during the first frosts.
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Development

In laboratory experiments, orange sulphurs took 31 days to mature from eggs to adults. Scott (1984) reports that the third and fourth stage larvae hibernate, while Opler (1984) states that orange sulphurs overwinter as crysales’.

Development - Life Cycle: metamorphosis ; diapause

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Life Expectancy

Lifespan/Longevity

Captive adult females have been found to live up to 39 days. In Virginia during a mark-release-recapture study, wild adult females had a lifespan of 14 days, males 25. If they overwinter, their entire lifespan may be almost a year.

Range lifespan

Status: captivity:
1 (high) years.

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Reproduction

Males spend their days patrolling their territories daily seeking females. They recognize the females visually, focusing on the coloration of the underside of the hindwing. The males are repelled by ultraviolet reflection on other males’ wings. Females appear not to care about the coloration of the males but ultraviolet reflection must be present, which helps reduce hybridization with yellow sulphurs Colias philodice that lack the reflection.

Female orange sulphurs begin to lay eggs when they have been adults for several days. In the lab they can lay up to 700 eggs. The eggs are laid singly in the middle of the upper surface of the host plant'’s leaf.

Breeding interval: Orange sulphurs have several broods throughout the warm seasons.

Range eggs per season: 700 (high) .

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous

There is no parental care given by adult butterflies.

Parental Investment: no parental involvement

  • Opler, P., G. Krizek. 1984. Butterflies East of the Great Plains. Baltimore: John Hopkins University Press.
  • Scott, J. 1986. The Butterflies of North America. Stanford, CA: Stanford University Press.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Colias eurytheme

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 9
Specimens with Barcodes: 56
Species With Barcodes: 1
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Barcode data: Colias eurytheme

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 11 barcode sequences available from BOLD and GenBank.  Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.  See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

GATATTGGAACTTTATATTTTATTTTTGGTGTGTGAGCAGGAATAATTGGAACTTCTTTA---AGTTTATTAATTCGTACAGAATTAGGTAATCCTGGATCACTAATTGGAGAT---GATCAAATTTATAACACTATTGTTACAGCTCATGCCTTTATTATAATTTTTTTTATAGTTATGCCAATTATAATTGGAGGATTTGGTAATTGATTAATTCCTTTGATA---TTAGGAGCCCCTGATATAGCTTTCCCACGTATAAATAATATAAGATTTTGATTACTACCCCCATCATTAACTTTATTAATTTCTAGAAGTATTGTTGAAAACGGAGCAGGAACAGGATGAACAGTTTACCCCCCTCTTTCCTCTAATATTGCCCATAGAGGATCTTCTGTTGATTTA---GCTATTTTTTCTCTTCACCTTGCAGGAATTTCCTCTATCCTAGGAGCAATTAATTTTATTACAACAATTATCAATATACGAATTAATAATATGTCATTTGATCAAATACCTTTATTTGTGTGAGCAGTAGGAATTACTGCTTTATTATTATTATTATCATTACCAGTTTTAGCTGGT---GCAATTACTATATTATTAACTGATCGGAATTTAAATACCTCTTTTTTTGACCCTGCTGGGGGAGGAGACCCAATTCTTTATCAACATTTATTTTGATTTTTTGGGCACCCCGAAGTATATATTCTAATTTTACCAGGATTTGGTATAATTTCACATATCATTTCTCAAGAAAGAGGGAAAAAA---GAAACTTTTGGATCTTTAGGAATAATTTATGCTATAATAGCAATTGGTTTATTAGGATTTATTGTATGAGCTCATCATATATTTACAGTTGGAATAGATATTGATACTCGAGCTTATTTCACCTCAGCAACTATAATTATTGCTGTACCTACAGGTATTAAAATTTTTAGTTGATTA---GCAACATTATATGGTACA---CAAATTAACTATAGTCCTTCTATATTATGAAGATTAGGATTTGTATTTTTATTTACTGTAGGGGGATTAACAGGGGTAATTTTAGCTAATTCATCTATTGATATTATTCTTCATGATACTTATTATGTTGTAGCACATTTCCATTATGTT---TTATCAATAGGAGCTGTATTTGCAATTTTAGGAGGATTTATTCATTGATACCCTCTATTTACAGGATTAATATTAAATCCATTTTATCTTAAAATTCAATTTATTACTATATTTATTGGAGTAAATTTAACTTTTTTTCCTCAACATTTTTTAGGTTTAGCTGGAATACCTCGT---CGATATTCAGATTACCCAGATAATTATCTT---TCTTGAAATATCATTTCATCGTTAGGATCTTATATTTCTTTAATTGGAACAATCATAATAATAATAATTATTTGAGAATCTATAATTAATAAACATTTTATT---ATTTTTTCAATAAATATACCTTCTTCT
-- end --

Download FASTA File
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Conservation

Conservation Status

Not of concern.
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National NatureServe Conservation Status

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N5 - Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: Widespread in lower 48 states and an abundant summer resident into Canada. Species has increased its range by about 40% since 1930. An occasional economic pest of alfalfa fields. This is one of the most common butterflies in the world and quite possibly the most common native butterfly in North America, and it increases each year as exotic legumes that the larvae do well on increase in the east and on top of that global warming can only increase its winter range, from which it colonizes much of the continent between June and September. The species is also somewhat migratory in the fall (Scott, 1986; Schweitzer, 2006) and part of the population emigrates from areas where it cannot survive the winter. Adults move north again in June.

Environmental Specificity: Broad. Generalist or community with all key requirements common.

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Orange sulphurs are stable rangewide.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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Global Short Term Trend: Increase of 10 to >25%

Global Long Term Trend: Increase of >25%

Comments: Has probably increased 100% or much more since 1800.

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Threats

Degree of Threat: Low

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Management

Global Protection: Unknown whether any occurrences are appropriately protected and managed

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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Orange sulphur caterpillars can be serious pests on alfalfa crops.

Negative Impacts: crop pest

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Economic Importance for Humans: Positive

Orange sulphurs provide enjoyment of people interested in butterfly watching.

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Wikipedia

Colias eurytheme

The Orange Sulphur (Colias eurytheme), also known as the Alfalfa Butterfly and in its larval stage as 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, but is absent from the central and southeastern USA.

Other members of this lineage including the Common or Clouded Sulphur (C. philodice) and Colias eriphyle and Colias vitabunda that 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 good species.[1]

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.

Distribution[edit]

Colias eurytheme butterflies can be found from southern Mexico to almost all throughout North America. Historically, they were distributed primarily in the Nearctic regions of the west, but were displaced to the east by logging and alfalfa field planting.[2]

Appearance[edit]

Wing Pattern[edit]

Male Colias eurytheme hind wings demonstrate an ultraviolet reflectance pattern while female Colias eurytheme hind wings 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 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.[3] 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.[4]

Genetic Inheritance[edit]

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 Colias 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.[5]

Reproduction[edit]

Reproductive behavior[edit]

Unlike that of many other butterfly species, the courtship of the Colias 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.[6]

These butterflies exhibit a polyandrous mating system. Upon mating, male Colias 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 remate once every 4 to 6 days in summer, and mate a lifetime total of up to 4 times.[7]

Sexual selection[edit]

Male Colias 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 sulfur butterflies Colias eurytheme and C. philodice.[8] The pheromone, located on the dorsal surface of the hindwing, consists of cuticular hydrocarbons n-heptacosane (C27), 13-methylheptacosane (13-MeC27), and possibly n-noncosane (C29).[9] In addition, wing scales located on the dorsal wing surfaces in male Colias eurytheme contain ridges with lamellae that produce iridescent ultraviolet reflectance via thin-film interference.[3]

Colias 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 hind wings. 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.[6]

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.[6] 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.[8]

Due to the widespread cultivation of the alfalfa, the host plant for Colias eurytheme and C. philodice, the species was able to expand their ranges across most of North America. These two species of sulfur butterflies have retained a large degree of genetic compatibility that allows them the produce viable and fertile offspring.[10] As a result of the recent sympatry and possible hybridization between these two species of sulfur butterflies, numerous studies have been conducted on intraspecific and interspecific mating.[11] 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 Colias eurytheme males.[10] Therefore, it was suggested that Colias eurytheme and C. philodice do not randomly mate with each other. Instead, mating was found to be positively assertive and mostly conspecific.[11]

Sexual selection theory[edit]

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.[12] 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.[7] 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.[8] 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.[13]

Gallery[edit]

References[edit]

  1. ^ 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)
  2. ^ Barton, Barb. "Colias eurytheme". Animal Diversity Web. University of Michigan Museum of Zoology. Retrieved 10 October 2013. 
  3. ^ a b 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.
  4. ^ 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.
  5. ^ 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.
  6. ^ a b c 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.
  7. ^ a b 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. 
  8. ^ a b c 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.
  9. ^ Sappington, T. W. (1990) Disruptive Sexual Selection in Colias Eurytheme Butterflies. Proceedings of the National Academy of Sciences 87(16):6132-5.
  10. ^ a b 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.
  11. ^ a b 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.
  12. ^ Rutowski, Ronald L. (1985) Evidence for Mate Choice in a Sulphur Butterfly (Colias eurytheme). Zeitschrift Für Tierpsychologie 70(2):103-14.
  13. ^ 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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Comments: Hybridizes with C. philodice, but not close to randomly. Great normal variation. Spring forms with reduced orange are not hybrids southward. Spring form adults where the January mean is more than a degree below freezing are generally male hybrids most years.

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