Parnassius smintheus

Males of this species often have poor visual discrimination, so they investigate all flying or sitting objects of the approximate size and color of the females of their species. They primarily identify females by their light color, so they are as likely to chase small blue lycaenids as they are to chase large white butterflies, but tend to ignore darker butterflies like fritillaries.[3] The males emerge from their pupae before females do in order to patrol for newly emerged females over a large area.[5] They patrol by flying over large areas then investigating any resting or flying females after spotting them. Males fly long distances continuously in search of females, stopping only to bask or feed.[3]

Males mate with young females, preferring females that have not yet expanded their wings. The females only mate once, lay eggs only once, and mature their eggs throughout their life, so the youngest females have the greatest potential fitness, and are selected for by males. There is also evidence that females emit a pheromone produced during the pupal stage, but its effect diminishes with time. The window of opportunity for a female to mate appears to be rather short, limited to only about three days.[5] The females of this species are not guaranteed to mate, and it has been estimated that around 14% of females never mate. Female mating success is correlated with population size, where reduced female mating success was observed at low population densities, and greater success at higher population densities. However, other studies have suggested that female mating success decreases as density increases, possibly due to male-male competition for mating.[9]

Mating plug in female's abdomen

The species has no courtship. Rather, males simply dive on a female upon finding her and force her down to the ground if she is not already there, and forcibly attempt to mate. This is successful if she has not already mated, but usually unsuccessful if she has already mated, due to a mating plug, a small waxy plug deposited onto a female's abdomen by a male to prevent future copulation. Unsuccessful matings may last an hour or more before the female escapes. If a female is unreceptive to mating, she may close up her wings tightly when a male approaches in an attempt to avoid being seen.[3]

After copulation, the male deposits a waxy genital plug on the tip of the female's abdomen, a mating plug, to prevent the female from mating again.[3] It contains sperm and nutrients for the female.[10] This ensures that the male is the only one to fertilize the female's eggs. Occasionally, the mating plug is not properly deposited, which means a mated female may sometimes present with no plug, or a mated male may present with a plug retained in his claspers.[3] However, no difference has been shown in the number of eggs laid by females with and females without a mating plug, indicating that male-donated nutrients play a negligible role in female fecundity. This suggests that in this species, the plug's primary role is to prevent the female butterfly from mating multiple times.[11]

Oviposition, or the laying of eggs occurs around the host plant Sedum lanceolatum, but not on the host plant itself. Females tend to search for good-quality meadows that have S. lanceolatum, though high host plant density is not required. They will retain their eggs and travel greater distances to find suitable meadows before oviposition. It is likely that there is some chemical or physical cue from the larval host plant that stimulates the females to oviposit near them. It is not as well understood why oviposition occurs off of the host plant, but researchers have suggested that it may be to avoid exposure of eggs to predators associated with the host plant.[12]

The eggs are white and round, but flattened at the top and bottom. They exhibit a pebbled surface, while the micropylar area, or the area where the sperm fertilized the egg, is brown and sunken.[3] The surface of the egg is generally more sculptured compared to the eggs of other butterflies, likely due to the thick chorion that has evolved to protect the overwintering egg from predators, parasitoids, and adverse environmental conditions.[13] Around 75 eggs are produced per female.[9] However, the eggshells are not consumed upon hatching, despite its high nutrient content.[14] The eggs are generally laid on the underside of flower heads, leaves, sticks, stones, moss, clumps of dirt, and sometimes on the larval foodplant. The female may or may not oviposit on the foodplant itself, and may often lay her eggs over a meter away from the plant. The embryo develops into first instar larva within a month of oviposition but the egg does not hatch until the snow melts the next spring.[3]

The larva completes five instars before pupating, developing over a period of around 10 to 12 weeks.[14][2] The first instar larva has a black body with many hairs, and a dull black head.[3] As the larva feeds on its host plant S. lanceolatum, it sequesters in its body sarmentosin from the plant, causing the larva to become distasteful to predators. Conspicuous yellow markings appear on the body after the second instar, warning off predators, as the larva stores higher levels of sarmentosin.[15] The fifth instar larva is black, with many short fine black hairs littered over its body. It sports two lateral and two dorsal rows of bright yellow spots. It has small and pale yellow vestigial osmeteria, a special organ just behind the head that resemble feelers. Since they may not always hatch close to their foodplant, the larvae have a fast, directional search pattern. They tend to feed rapidly on their larval host plant, then move to a basking location on open ground up to a few meters away for thermoregulation.[3] If disturbed, the larvae twitch violently, then drop to the ground to seek cover, and may discharge a bad-smelling brown chemical.[14]

The pupae are dark yellow brown to red brown and are formed in the leaf litter, gravel, or rocks lying near the soil surface.[3] The larvae do not spin a cocoon but rather gather some silk around them simply to attach themselves to the surroundings.[12] When ready to emerge, newly developed butterflies have two tiny hooks to assist in emerging from where they were pupating.[10]

Adult

Adults are generally translucent yellowish white with black, grey, and red markings, though they vary greatly in color.[3] They have a wingspan ranging from 2 to 3 inches. The dorsal forewing exhibits a marginal grey band and a weakly developed pale grey submarginal band. The margin of the forewing usually has small triangles of black at each vein running through the wing, as well as some grey markings. The ventral hindwing may also have marginal and submarginal grey markings. The fringes of the wings are typically black at then ends of the veins. The hairs and scales on the heads, legs, and ventral abdomen are usually yellowish. Both sexes usually have bright red spots on the hindwings and forewings. Populations residing at higher elevations typically have darker females, and both males and females are smaller at low elevations.[3] Adult butterflies of this species feed predominantly on the nectar of yellow-flowered species including Potentilla fruticosa, Solidago multiradiata, Senecio canus, and S. lanceolatum.[4]

The larvae sequester sarmentosin from their food plant to gain protection from predators; an individual butterfly contains around 460 ɥg of sarmentosin. The closely related P. apollo also sequesters sarmentosin, but the concentrations were found to be nearly three times higher in P. smintheus adults. The concentration of sarmentosin is highest in the wings of the butterfly, which helps fend off predators since the wings and its scales are the first thing to come in contact with any predators.[15] As a result of sequestering the compound, the butterflies have a strong odor and excrete brown fluid from their anus that smells like their body when attacked. Their body fluid is a nasal irritant to humans.[3] The exact defense mechanism of sarmentosin is not known, but it may be a very bitter compound, making predators unlikely to prey on them again.[15] Their white wings with black and red markings warn of their unpalatability to birds, while their odor serves to warn rodents off. Males most likely have more distasteful compounds in them than females, because chipmunks tend to eat more females than males of this species.[3]