1053:, where its numbers have declined to the point that based on IUCN criterion, it has reached endangered species status. The causes of this population decline are not fully determined, but there are several possible factors. Since the common brimstone is univoltine, it may have difficulties adapting to changing environmental conditions compared to species that have multiple generations a year. For example, there has been a decrease in suitable overwintering environments for the butterflies, with open woodland decreasing in favour of more urban areas. Nitrogen pollution, declining nectar supplies, and rapid ecological changes have also been suggested as other hypothetical factors. Concerns have been raised about the possible future increase of this population decline, but the butterfly mostly does not appear to be a conservation concern due to its widespread and common geographic presence.
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develop, and pupate, newly hatched adult butterflies emerge and disperse locally into both woodlands and wetlands to overwinter. Butterflies travel to the woodlands for overwintering, and no mating appears to occur within these habitats. Overwintering also occurs in the wetlands, where the host plant alder buckthorn is abundant. After emerging from overwintering, adult brimstones that were previously in the wetlands are joined by those that hibernated in woodlands, and the population breeds and lays eggs.
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iridescence is seen exclusively under ultraviolet light, since females absorb light on the ultraviolet spectra. The presence of exclusively leucopterin in female wings explains the lack of iridescence in female common brimstones, since leucopterin absorbs only in the ultraviolet range. Therefore, the wings do not reflect and consequently do not scatter any ultraviolet light like male wings do.
936:. This has been possibly attributed to several factors, such as the greater abundance and quality of resources in areas with these environmental conditions. Other possibilities include a better ability to assimilate resources as an indication of male mate quality. Because ultraviolet coloration is energetically expensive to develop, it could signal high male quality.
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activities. Downhill migration is influenced by the need for larval resources such as host plants during breeding seasons - the butterflies travel to lower elevations in search for regions containing these plants, with adults commonly returning to the areas where they had been bred due to their long lifespan.
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colouration, meaning they match the colour of their habitats. Larvae are so difficult to see due to this colouration that they can remain in the open undetected. When not eating, the caterpillars remain still in a position alongside the midrib of leaves, making them even more difficult to spot. Adult
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is approximately 50 days. However, the adult brimstone spends a large portion of its life in an overwintering state. The brimstone is highly mobile, feeding and travelling to regions ideal for hibernation during the late summer and fall, and returning to regions ideal for mating and egg-laying during
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mating patterns, males may emerge earlier to increase the number of mating chances and therefore reproductive success, as older males have had more time to develop and therefore have a greater advantage. In contrast, females emerge late due to the late seasonal development of host plants such as the
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was extracted from the white wings of females. The difference in wing pigmentation contributes to the distinct iridescence patterns of males and females. Iridescence occurs due to visualised changes in coloration from the scattering of ultraviolet light. A male-only pattern of coloration due to this
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The adult common brimstone overwinters for seven months, remaining hidden and motionless throughout its hibernation. While both sexes have similar egg to adult development times, they differ in the times that they reach sexual maturity. The reproductive development of males begins just after pupal
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responses whether they were presented in natural or synthetic mixes of floral compounds. Additionally, these two compounds are present in the largest quantities in the nectar plants utilised by the brimstone, indicating that scent detection could be important for detecting food sources. This would
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migrations of the common brimstone. Uphill migration is potentially influenced by habitat limitations, such as a lack of the forest cover that is required during overwintering. Brimstones travel to higher elevations for greater forest cover and reduced exposure to higher temperatures during their
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are attracted to damaged plants through chemical or visual signals, less damage leads to greater offspring survival since eggs are less likely to be detected. Plants exposed to both sunlight and the open lead to reduced chances of predation and parasitism as well, and are more accessible to adult
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Adult common brimstones lay eggs singly on the underside of buckthorn leaves. The eggs are around 1.3 mm tall, and are spindle-shaped in appearance. The eggs change colour over time, initially having a greenish-white colouration, then progressively darker shades of yellow, and finally brown
526:, meaning that it lays one generation of eggs each year. There are several ideal characteristics of the particular host plants chosen for oviposition. Adult brimstones lay eggs on the underside of the leaves of the two species of host plants, where they are less conspicuous. The high mobility of
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Adults emerge during the summer, from June to August, and continue to feed until
September. The common brimstone hibernates for the next seven months of winter, remaining inactive until April, where they then emerge and proceed to reproduce and lay eggs. Adult brimstones are highly abundant for
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The common brimstone undergoes some regional migration between hibernation and breeding areas throughout the year, as seen in the different chemical composition of butterflies across varying seasons and regions. In general, there is movement towards wetlands to reproduce. After the eggs hatch,
404:. The high mobility of this butterfly allows it to search widely for new host plant locations and expand its range. While the geographic distribution of the adult is larger than that of its host plant, its range is nevertheless limited by the presence of host plants due to the needs of its
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In males, iridescence is indicated in that the wing pattern appears to visually change depending on the position of the ultraviolet light shone onto the wing. At some angles, a male pattern is seen, while at other angles, a female lack of pattern is seen. This is referred to as the
354:. In spring when their host plants have developed, they return to the wetlands to breed and lay eggs. Both the larval and adult forms of the common brimstone have protective coloration and behaviour that decreases their chances of being recognised and subsequently preyed upon.
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emergence, and continues during hibernation, which indicates that males may not be able to reproduce until after overwintering. For females, eggs remain undeveloped as the butterflies overwinter, and no reproductive development occurs until after emergence from hibernation.
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in females throughout the mating season. Pairs are formed after the butterflies have engaged in a dalliance flight for a period of time. When a pair settles to mate, they do not take flight during copulation and remain paired for a long time of up to forty-eight hours.
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effect". This demonstrates that the pattern appears to be optical, rather than pigmental, as the effect is only seen at certain angles and distances of light and changes with positions. If it were pigmental, these changes would not cause differences in iridescence.
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The common brimstone is one of the longest-living butterflies, with a life expectancy ranging from 10 months to a year. Due to its hibernation and life cycle, it has one generation per year. Development from the laid egg to the emergence of the
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The sexes also differ in times of emergence after overwintering. Emergence is correlated with temperature and hours of sunlight; a certain amount of both is necessary for the butterfly to emerge from hibernation and therefore influences when
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brimstones are leaf-mimics, as they share similarities in shape, colour, and pattern to leaves. This allows them to blend in with their surroundings during vulnerable times like diapause (hibernation). When picked up, the butterflies become
837:, possibly due to the wide distribution of the butterfly and the host plants in its habitats. The broad presence of its host allows the wasps to be host-specific. The wasps are primarily associated with the presence of the food plant
423:. The common brimstone prefers laying eggs on younger host plants with late bud-bursts that are isolated from other plants in the area and exposed to both open space and sun. During the winter, adult brimstones travel to woodlands to
1093:. Swaay, Chris van., European Commission. Directorate-General for Environment. International Union for Conservation of Nature and Natural Resources. Blue List Programme. Luxembourg: Publications Office of the European Union. 2010.
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components. These differences cause light to scatter in different ways, leading to the different colours. In the common brimstone, wing scales scatter light incoherently due to ovoid-shaped structures called beads that contain
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across its length. When they first hatch, they move to the top side of the leaves and eat them, leaving characteristic hole patterns in their host plants. During the day, they feed and then rest in the open, lying still on the
484:
Unlike their larval forms, which are specialised for particular host plants, adult brimstones are not specialised nectar feeders. The common brimstone heavily feeds on the nectar of several flowering species including knapweed
1145:
Pecháček, Pavel; Stella, David; Keil, Petr; Kleisner, Karel (2014-12-01). "Environmental effects on the shape variation of male ultraviolet patterns in the
Brimstone butterfly (Gonepteryx rhamni, Pieridae, Lepidoptera)".
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occurs over approximately two weeks. The pupae are 22.2-23.8 mm in length and have the appearance of a curled leaf, with pointed ends and bulges in the middle. The pupae are secured to stems and leaves using silk; a
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enables the butterflies to find even the most isolated host plants in an area, which are more ideal for their offspring. Eggs are more likely to be deposited on outlying plants, leading to reduced vulnerability as fewer
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butterflies. Larvae can also benefit from decreased host plant defences; juvenile plants and plants with late bud-bursts produce fewer toxic defence chemicals, as resources are directed more towards plant growth.
435:. The common brimstone has an appearance that is highly similar to the leaves of these plants, so during hibernation it can remain hidden. In other seasons, habitat selection is also affected by the abundance of
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are attracted to these plants. Another factor is damage; undamaged plants indicate the absence of other eggs, as brimstone larvae leave holes in the leaves of the plants on which they feed. Since predators and
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hook attaches to a silk padding, and a length of silk secures the pupae around its middle. The pupae have a primarily green colouration, but right before adult emergence for males, the wing areas turn yellow.
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plants as host plants for its larvae; this influences its geographic range and distribution, as these plants are commonly found in wetlands. The adult brimstone travels to woodland areas to spend seven months
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does not appear to have a threatened conservation status according to IUCN standards. However, the butterfly has experienced significant population and distribution reduction in areas such as the
576:, initially having a length of 1.7 mm in the first instar and reaching up to 34.9 mm in length when fully grown. The caterpillars have a green colouration with white hairs and dark
1841:
Lozan, Aurel; Spitzer, Karel; Jaroš, Josef (2012-06-01). "Isolated peat bog habitats and their food connections: parasitoids (Hymenoptera: Ichneumonoidea) and their lepidopteran hosts".
1991:
Wiklund, Christer; Forsberg, Johan (1991). "Sexual Size
Dimorphism in Relation to Female Polygamy and Protandry in Butterflies: A Comparative Study of Swedish Pieridae and Satyridae".
509:) in April and May and have been recorded gathering nectar from many other species of flowers. Adult food plant availability is another factor that is important for habitat selection.
598:
1340:
Wiklund, Christer; Lindfors, Virpi; Forsberg, Johan (1996). "Early Male
Emergence and Reproductive Phenology of the Adult Overwintering Butterfly Gonepteryx rhamni in Sweden".
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in adult butterflies, as odour could act as a cue for finding and distinguishing nectar plants, allowing more energy to be utilised for other activities such as reproduction.
2190:
Van Dyck, Hans; Van Strien, Arco J.; Maes, Dirk; Van Swaay, Chris a. M. (2009-08-01). "Declines in Common, Widespread
Butterflies in a Landscape under Intense Human Use".
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yellow wing colouration and females having a greenish-white wing colouration. Additionally, males have iridescent dorsal wings that change in colour and appearance under
2095:
Andersson, Susanna (2003-03-01). "Antennal responses to floral scents in the butterflies
Inachis io, Aglais urticae (Nymphalidae), and Gonepteryx rhamni (Pieridae)".
2037:
Andersson, Susanna (2003-03-01). "Foraging responses in the butterflies
Inachis io, Aglais urticae (Nymphalidae), and Gonepteryx rhamni (Pieridae) to floral scents".
1541:
Dempster, J. P.; Lakhani, K. H.; Coward, P. A. (1986-02-01). "The use of chemical composition as a population marker in insects: a study of the
Brimstone butterfly".
924:
The structural coloration of the male dorsal wings is affected by environmental factors. There is an increase in ultraviolet coloration coverage with increasing
1945:
Nekrutenko, Yuri P. (January 1965). "'Gynandromorphic Effect' and the
Optical Nature of Hidden Wing-pattern in Gonepteryx rhamni; L. (Lepidoptera. Pieridae)".
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ends. Males emerge earlier than females, as they are more willing to fly in lower temperatures than females. Since the common brimstone most closely follows
476:. This influences the distribution of the adult brimstone, as the presence of these two buckthorn species is necessary for the survival of their offspring.
944:
After the common brimstone emerges from hibernation, it travels towards habitats that contain larval host plants and mates. The brimstone is primarily
1785:
Bibby, Tina J. (1983). "Oviposition by the
Brimstone Butterfly, Gonepteryx Rhamni (L.)(Lepidoptera: Pieridae) in Monks Wood, Cambridgeshire in 1982".
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Through chemical extraction and analysis, two possible pigments have been identified that may contribute to the common brimstone's wing coloration.
742:
727:
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1643:
Jennersten, Ola (1980). "Nectar source plant selection and distribution pattern in an autumn population of Gonepteryx rhamni (Lep. Pieridae)".
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Pollard, E.; Hall, M. L. (1980). "Possible movement of Gonepteryx rhamni (L.) (Lepidoptera: Pieridae) between hibernating and breeding areas".
712:
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1485:
McKay, H. V. (1991). "Egg-Laying Requirements of Woodland Butterflies; Brimstones (Gonepteryx rhamni) and Alder Buckthorn (Frangula alnus)".
908:
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1701:
810:. Both larvae and adult brimstones fall victim to predation and use means such as protective coloration and mimicry to avoid this fate.
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2599:
2447:
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Gutiérrez, David; Thomas, Chris D. (2000-05-01). "Marginal range expansion in a host-limited butterfly species Gonepteryx rhamni".
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also has a stronger reliance on visual indications such as colour compared with other butterfly species, which rely more on odour.
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alder buckthorn, since these plants are necessary for egg-laying. Female emergence is correlated with host plant development.
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2657:
1062:
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Fox-Wilson, G. (1940-03-01). "Phenological Observations on the Brimstone Butterfly, Gonepteryx Rhamni (linn.) (lepid.)".
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compounds oxoisophoroneoxide, oxoisophorone, and dihydrooxoisophorone, as these compounds elicited some of the strongest
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several months after their emergence from overwintering. The common brimstone has sexual dichromism, with males having a
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while females have greenish-white wings and are not iridescent. This iridescence is affected by environmental factors.
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occurs in the presence of certain compounds. Research suggests that there are antennal olfactory receptors for
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The common brimstone appears to have an innate preference for certain colours in nectar plants – red and blue
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The common brimstone uses various environments for different stages of its life cycle. The butterfly inhabits
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1702:"Climate conditions and resource availability drive return elevational migrations in a single-brooded insect"
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light and scatter longer wavelengths outside of the pigment absorption spectrum, such as light in the
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is responsible for the sulphur yellow colour of the male wings since it absorbs in the violet range.
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A Natural History of the British Lepidoptera: A Text-Book for Students and Collectors: by J. W. Tutt
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335:. Its wing span size is 60–74 mm (2.4–2.9 in). It should not be confused with the
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response to the floral scent compounds of nectar plants, where neural activity in antennal
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The environmental conditions of a particular year also affect migration, as seen in the
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and hide their legs from view in order to decrease their chances of being recognised.
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Brakefield, Paul M.; Shreeve, Tim G. (1992). "Avoidance, concealment, and defence".
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503:). However, brimstones have also been observed feeding on the nectar of coltsfoot (
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light, while females do not. Both males and females have orange spots in the
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It was first described and published in Linnaeus's book, the 10th edition of
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of each wing, pink head and antennae, and a thorax covered in white hair.
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of leaves, where their colouration makes them difficult to distinguish.
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2020:
1886:"Wing coloration and pigment gradients in scales of pierid butterflies"
1825:. Vol. 6. London: Swan Sonnenschein & Co. pp. 12, 45, 75.
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833:. These two species of parasitoid wasps are completely specialised for
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1459:"Gonepteryx rhamni Linnaeus, 1758 - Common Brimstone | Butterfly"
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Variation in coloration of Lepidoptera wings is caused by different
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during mating and breeding season, as they provide ideal areas for
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The common brimstone has two recorded species of parasites: the
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Larval brimstones appear to feed on only two plant sources: the
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Wijnen, B.; Leertouwer, H. L.; Stavenga, D. G. (2007-12-01).
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Frohawk, F. W. (1940). "Food-plants of Gonepteryx rhamni".
1625:. Vol. 1. London: Hutchinson & Co. pp. 58–63.
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are common in heavily used nectar sources in some regions.
948:, as demonstrated by the presence of usually only a single
807:
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384:, the colour which matches the colour of the male's wings.
2242:
The Millennium Atlas of Butterflies of Britain and Ireland
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Proceedings of the Royal Entomological Society of London A
1402:"Colors and pterin pigmentation of pierid butterfly wings"
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The common brimstone can be commonly found throughout the
874:
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Jim Asher, Martin Warren & Richard Fox, ed. (2001).
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1986:
1984:
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Giraldo, Marco A.; Stavenga, Doekele G. (2008-03-01).
327:. Across much of its range, it is the only species of
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Tuft, James William (November 1905 – December 1906).
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1981:
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The common buckthorn, one of the larval host plants.
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in its wing coloration: males have yellow wings and
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zone and is commonly found across Europe, Asia, and
857:Both the larval and adult common brimstone exhibit
1700:Gutiérrez, David; Wilson, Robert J. (2014-07-01).
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892:. Due to these pigments, the beads absorb short
572:The larvae of the common brimstone undergo five
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331:, and is therefore simply known locally as the
1310:. London: Oxford University Press. p. 98.
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419:due to an abundance of host plants like the
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1944:
1921:11370/98cb9aa4-47c9-4849-af82-d4c4fcc79ec1
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1623:The Natural History of British Butterflies
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853:Larva lying alongside the midrib of a leaf
396:. Individuals have been seen from western
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642:Mating pair (left: male; right: female)
439:as a food source for adult brimstones.
345:The brimstone relies on two species of
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1308:The Ecology of Butterflies in Britain
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1063:List of butterflies of Great Britain
845:Protective colouration and behaviour
2628:IUCN Red List least concern species
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1935:
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13:
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2160:10.1111/j.1365-3032.1940.tb00570.x
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1563:10.1111/j.1365-2311.1986.tb00279.x
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802:is preyed upon by many species of
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2212:10.1111/j.1523-1739.2009.01175.x
1232:10.1046/j.1365-2311.2000.00241.x
1091:European red list of Butterflies
798:Like most woodland Lepidoptera,
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357:The adult common brimstone has
1843:Journal of Insect Conservation
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1429:10.1016/j.jinsphys.2007.06.016
666:Caterpillar on alder buckthorn
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1:
2653:Butterflies described in 1758
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1003:contribute to more efficient
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380:Brimstone is an old name for
1409:Journal of Insect Physiology
982:The common brimstone has an
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2658:Taxa named by Carl Linnaeus
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870:Genetics of colour patterns
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1487:Journal of Applied Ecology
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319:. It lives throughout the
59:Female at Parsonage Moor,
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1913:10.1016/j.asd.2007.09.003
1855:10.1007/s10841-011-9425-4
1729:10.1007/s00442-014-2952-4
1168:10.1007/s00114-014-1244-5
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99:Scientific classification
97:
75:
66:
58:
49:
37:
28:
23:
522:The common brimstone is
479:
388:Distribution and habitat
340:Opisthograptis luteolata
2246:Oxford University Press
1621:Frohawk, F. W. (1924).
588:
1787:Entomologist's Gazette
1645:Entomologisk Tidskrift
1274:Entomologist's Gazette
1121:: CS1 maint: others (
1020:
1019:Hibernating adult male
854:
558:
452:
2648:Butterflies of Africa
2643:Butterflies of Europe
2117:10.1007/s000490300001
2059:10.1007/s000490300000
1543:Ecological Entomology
1212:Ecological Entomology
1018:
852:
450:
307:, commonly named the
2435:Fauna Europaea (new)
2278:at Wikimedia Commons
2192:Conservation Biology
1000:electrophysiological
824:Cotesia gonopterygis
474:Rhamnus carthartica)
16:Species of butterfly
2638:Butterflies of Asia
2204:2009ConBi..23..957V
2109:2003Checo..13...13A
2051:2003Checo..13....1A
2005:1991Oikos..60..373W
1959:1965Natur.205..417N
1905:2008ArtSD..37..118G
1721:2014Oecol.175..861G
1555:1986EcoEn..11...51D
1499:1991JApEc..28..731M
1421:2007JInsP..53.1206W
1354:1996Oikos..75..227W
1224:2000EcoEn..25..165G
1160:2014NW....101.1055P
1148:Naturwissenschaften
988:olfactory receptors
69:Conservation status
1021:
992:phenylacetaldehyde
900:wavelength range.
855:
453:
251:G. r. miljanowskii
246:G. r. meridionalis
2615:
2614:
2559:Open Tree of Life
2338:Gonepteryx rhamni
2308:Gonepteryx rhamni
2300:Taxon identifiers
2288:Gonepteryx rhamni
2276:Gonepteryx rhamni
2274:Media related to
2255:978-0-19-850565-5
1953:(4969): 417–418.
1415:(12): 1206–1217.
1154:(12): 1055–1063.
932:, and decreasing
764:Gonepteryx rhamni
749:Gonepteryx rhamni
734:Gonepteryx rhamni
719:Gonepteryx rhamni
564:before hatching.
506:Tussilago farfara
500:Succisa pratensis
359:sexual dimorphism
304:Gonepteryx rhamni
300:
299:
294:
261:G. r. tianshanica
208:Gonepteryx rhamni
92:
24:Common brimstone
2665:
2608:
2607:
2595:
2594:
2593:
2567:
2566:
2554:
2553:
2541:
2540:
2528:
2527:
2525:NHMSYS0000516380
2515:
2514:
2505:
2504:
2495:
2494:
2482:
2481:
2469:
2468:
2456:
2455:
2443:
2442:
2430:
2429:
2417:
2416:
2404:
2403:
2391:
2390:
2378:
2377:
2365:
2364:
2352:
2351:
2342:
2341:
2340:
2327:
2326:
2325:
2295:
2294:
2286:Data related to
2285:
2273:
2259:
2232:
2231:
2187:
2172:
2171:
2143:
2137:
2136:
2092:
2079:
2078:
2034:
2025:
2024:
1988:
1979:
1978:
1967:10.1038/205417a0
1942:
1933:
1932:
1890:
1881:
1875:
1874:
1838:
1827:
1826:
1816:
1803:
1802:
1782:
1767:
1766:
1740:
1706:
1697:
1680:
1679:
1672:The Entomologist
1667:
1661:
1660:
1640:
1627:
1626:
1618:
1583:
1582:
1538:
1519:
1518:
1482:
1463:
1462:
1455:
1449:
1448:
1406:
1397:
1374:
1373:
1337:
1312:
1311:
1303:
1290:
1289:
1269:
1252:
1251:
1207:
1196:
1195:
1142:
1127:
1126:
1120:
1112:
1087:
760:
745:
730:
715:
699:
687:
675:
663:
651:
639:
494:Knautia arvensis
491:) and scabious (
470:common buckthorn
309:common brimstone
286:
241:G. r. kurdistana
210:
107:
106:
86:
81:
80:
54:
38:Male in flight,
33:
21:
20:
2673:
2672:
2668:
2667:
2666:
2664:
2663:
2662:
2618:
2617:
2616:
2611:
2603:
2598:
2589:
2588:
2583:
2570:
2562:
2557:
2549:
2546:Observation.org
2544:
2536:
2531:
2523:
2518:
2510:
2508:
2500:
2498:
2490:
2485:
2477:
2472:
2464:
2459:
2451:
2446:
2438:
2433:
2425:
2420:
2412:
2407:
2399:
2394:
2386:
2381:
2373:
2368:
2360:
2355:
2347:
2345:
2336:
2335:
2330:
2321:
2320:
2315:
2302:
2266:
2256:
2236:
2235:
2188:
2175:
2144:
2140:
2093:
2082:
2035:
2028:
2013:10.2307/3545080
1989:
1982:
1943:
1936:
1888:
1882:
1878:
1839:
1830:
1817:
1806:
1783:
1770:
1704:
1698:
1683:
1668:
1664:
1641:
1630:
1619:
1586:
1539:
1522:
1507:10.2307/2404579
1483:
1466:
1457:
1456:
1452:
1404:
1398:
1377:
1362:10.2307/3546246
1338:
1315:
1304:
1293:
1270:
1255:
1208:
1199:
1143:
1130:
1114:
1113:
1101:
1089:
1088:
1081:
1076:
1059:
1043:
1013:
980:
964:
959:
942:
918:gynandromorphic
877:
872:
847:
830:Cotesia risilis
816:
796:
791:
774:
767:
761:
752:
746:
737:
731:
722:
716:
707:
706:
705:
704:
703:
700:
692:
691:
688:
680:
679:
676:
668:
667:
664:
656:
655:
652:
644:
643:
640:
631:
630:
608:
591:
570:
561:
547:
520:
515:
488:Centaurea jacea
482:
462:alder buckthorn
458:
445:
421:alder buckthorn
390:
375:Systema Naturae
371:
266:G. r. transiens
223:
212:
206:
193:
101:
93:
82:
78:
71:
17:
12:
11:
5:
2671:
2661:
2660:
2655:
2650:
2645:
2640:
2635:
2630:
2613:
2612:
2610:
2609:
2596:
2580:
2578:
2576:Papilio rhamni
2572:
2571:
2569:
2568:
2555:
2542:
2529:
2516:
2506:
2496:
2483:
2470:
2457:
2444:
2431:
2422:Fauna Europaea
2418:
2405:
2392:
2379:
2366:
2353:
2343:
2328:
2312:
2310:
2304:
2303:
2292:
2291:
2290:at Wikispecies
2279:
2265:
2264:External links
2262:
2261:
2260:
2254:
2234:
2233:
2198:(4): 957–965.
2173:
2154:(1–3): 13–16.
2138:
2080:
2026:
1999:(3): 373–381.
1980:
1934:
1899:(2): 118–128.
1876:
1849:(3): 391–397.
1828:
1804:
1768:
1715:(3): 861–873.
1681:
1662:
1628:
1584:
1520:
1493:(2): 731–743.
1464:
1450:
1375:
1348:(2): 227–240.
1313:
1291:
1253:
1218:(2): 165–170.
1197:
1128:
1099:
1078:
1077:
1075:
1072:
1071:
1070:
1068:Brimstone moth
1065:
1058:
1055:
1042:
1039:
1012:
1009:
979:
976:
968:inflorescences
963:
960:
958:
955:
941:
938:
876:
873:
871:
868:
846:
843:
839:Frangula alnus
815:
812:
795:
792:
790:
787:
773:
770:
769:
768:
762:
755:
753:
747:
740:
738:
732:
725:
723:
717:
710:
701:
694:
693:
689:
682:
681:
677:
670:
669:
665:
658:
657:
653:
646:
645:
641:
634:
633:
632:
628:
627:
626:
625:
621:discoidal cell
607:
604:
590:
587:
569:
566:
560:
557:
546:
543:
519:
516:
514:
511:
481:
478:
466:Frangula alnus
457:
454:
444:
443:Food resources
441:
389:
386:
370:
367:
337:brimstone moth
315:of the family
298:
297:
296:
295:
284:Papilio rhamni
278:
277:
271:
270:
269:
268:
263:
258:
253:
248:
243:
238:
230:
229:
225:
224:
213:
202:
201:
195:
194:
190:G. rhamni
187:
185:
181:
180:
173:
169:
168:
163:
159:
158:
153:
149:
148:
143:
139:
138:
133:
129:
128:
123:
119:
118:
113:
109:
108:
95:
94:
76:
73:
72:
67:
64:
63:
56:
55:
47:
46:
35:
34:
26:
25:
15:
9:
6:
4:
3:
2:
2670:
2659:
2656:
2654:
2651:
2649:
2646:
2644:
2641:
2639:
2636:
2634:
2631:
2629:
2626:
2625:
2623:
2606:
2601:
2597:
2592:
2586:
2582:
2581:
2579:
2577:
2573:
2565:
2560:
2556:
2552:
2547:
2543:
2539:
2534:
2530:
2526:
2521:
2517:
2513:
2507:
2503:
2497:
2493:
2488:
2484:
2480:
2475:
2471:
2467:
2462:
2458:
2454:
2449:
2445:
2441:
2436:
2432:
2428:
2423:
2419:
2415:
2410:
2406:
2402:
2397:
2393:
2389:
2384:
2380:
2376:
2371:
2367:
2363:
2358:
2354:
2350:
2344:
2339:
2333:
2329:
2324:
2318:
2314:
2313:
2311:
2309:
2305:
2301:
2296:
2289:
2284:
2280:
2277:
2272:
2268:
2267:
2257:
2251:
2247:
2243:
2238:
2237:
2229:
2225:
2221:
2217:
2213:
2209:
2205:
2201:
2197:
2193:
2186:
2184:
2182:
2180:
2178:
2169:
2165:
2161:
2157:
2153:
2149:
2142:
2134:
2130:
2126:
2122:
2118:
2114:
2110:
2106:
2102:
2098:
2091:
2089:
2087:
2085:
2076:
2072:
2068:
2064:
2060:
2056:
2052:
2048:
2044:
2040:
2033:
2031:
2022:
2018:
2014:
2010:
2006:
2002:
1998:
1994:
1987:
1985:
1976:
1972:
1968:
1964:
1960:
1956:
1952:
1948:
1941:
1939:
1930:
1926:
1922:
1918:
1914:
1910:
1906:
1902:
1898:
1894:
1887:
1880:
1872:
1868:
1864:
1860:
1856:
1852:
1848:
1844:
1837:
1835:
1833:
1824:
1823:
1815:
1813:
1811:
1809:
1800:
1796:
1792:
1788:
1781:
1779:
1777:
1775:
1773:
1764:
1760:
1756:
1752:
1748:
1744:
1739:
1734:
1730:
1726:
1722:
1718:
1714:
1710:
1703:
1696:
1694:
1692:
1690:
1688:
1686:
1677:
1673:
1666:
1658:
1654:
1650:
1646:
1639:
1637:
1635:
1633:
1624:
1617:
1615:
1613:
1611:
1609:
1607:
1605:
1603:
1601:
1599:
1597:
1595:
1593:
1591:
1589:
1580:
1576:
1572:
1568:
1564:
1560:
1556:
1552:
1548:
1544:
1537:
1535:
1533:
1531:
1529:
1527:
1525:
1516:
1512:
1508:
1504:
1500:
1496:
1492:
1488:
1481:
1479:
1477:
1475:
1473:
1471:
1469:
1460:
1454:
1446:
1442:
1438:
1434:
1430:
1426:
1422:
1418:
1414:
1410:
1403:
1396:
1394:
1392:
1390:
1388:
1386:
1384:
1382:
1380:
1371:
1367:
1363:
1359:
1355:
1351:
1347:
1343:
1336:
1334:
1332:
1330:
1328:
1326:
1324:
1322:
1320:
1318:
1309:
1302:
1300:
1298:
1296:
1287:
1283:
1279:
1275:
1268:
1266:
1264:
1262:
1260:
1258:
1249:
1245:
1241:
1237:
1233:
1229:
1225:
1221:
1217:
1213:
1206:
1204:
1202:
1193:
1189:
1185:
1181:
1177:
1173:
1169:
1165:
1161:
1157:
1153:
1149:
1141:
1139:
1137:
1135:
1133:
1124:
1118:
1110:
1106:
1102:
1100:9789279141515
1096:
1092:
1086:
1084:
1079:
1069:
1066:
1064:
1061:
1060:
1054:
1052:
1048:
1038:
1035:
1031:
1025:
1017:
1008:
1006:
1001:
997:
993:
989:
985:
975:
973:
969:
954:
951:
950:spermatophore
947:
937:
935:
931:
930:precipitation
928:, increasing
927:
922:
919:
913:
910:
906:
901:
899:
898:complementary
895:
891:
886:
882:
867:
865:
860:
851:
842:
840:
836:
832:
831:
826:
825:
821:
811:
809:
805:
801:
786:
783:
778:
765:
759:
754:
750:
744:
739:
735:
729:
724:
720:
714:
709:
708:
698:
686:
674:
662:
650:
638:
624:
622:
618:
614:
603:
600:
595:
586:
584:
579:
575:
565:
556:
553:
542:
539:
534:
529:
525:
513:Parental care
510:
508:
507:
502:
501:
496:
495:
490:
489:
477:
475:
471:
467:
463:
449:
440:
438:
434:
430:
426:
422:
418:
414:
409:
407:
403:
399:
395:
385:
383:
378:
376:
366:
364:
360:
355:
353:
352:overwintering
348:
343:
341:
338:
334:
330:
326:
322:
318:
314:
310:
306:
305:
293:
289:
285:
282:
281:
279:
276:
272:
267:
264:
262:
259:
257:
254:
252:
249:
247:
244:
242:
239:
237:
236:G. r. gravesi
234:
233:
231:
226:
221:
217:
211:
209:
203:
200:
199:Binomial name
196:
192:
191:
186:
183:
182:
179:
178:
174:
171:
170:
167:
164:
161:
160:
157:
154:
151:
150:
147:
144:
141:
140:
137:
134:
131:
130:
127:
124:
121:
120:
117:
114:
111:
110:
105:
100:
96:
90:
85:
84:Least Concern
74:
70:
65:
62:
57:
53:
48:
45:
41:
36:
32:
27:
22:
19:
2575:
2307:
2241:
2195:
2191:
2151:
2147:
2141:
2103:(1): 13–20.
2100:
2097:Chemoecology
2096:
2042:
2039:Chemoecology
2038:
1996:
1992:
1950:
1946:
1896:
1892:
1879:
1846:
1842:
1821:
1790:
1786:
1712:
1708:
1675:
1671:
1665:
1648:
1644:
1622:
1549:(1): 51–65.
1546:
1542:
1490:
1486:
1453:
1412:
1408:
1345:
1341:
1307:
1277:
1273:
1215:
1211:
1151:
1147:
1090:
1046:
1045:As of 2010,
1044:
1041:Conservation
1026:
1022:
981:
971:
965:
943:
923:
914:
905:Xanthopterin
902:
878:
856:
838:
834:
828:
822:
817:
799:
797:
779:
775:
763:
748:
733:
718:
702:Adult female
609:
592:
571:
562:
555:the spring.
548:
527:
521:
504:
498:
492:
486:
483:
473:
465:
459:
431:foliage and
410:
391:
379:
372:
356:
344:
339:
332:
325:North Africa
308:
303:
302:
301:
283:
265:
260:
256:G. r. rhamni
255:
250:
245:
240:
235:
207:
205:
189:
188:
176:
40:Dry Sandford
18:
2461:iNaturalist
2332:Wikispecies
2045:(1): 1–11.
1793:: 229–234.
1738:10871/16841
1651:: 109–114.
1280:: 217–220.
1051:Netherlands
926:temperature
909:Leucopterin
782:elevational
617:ultraviolet
568:Caterpillar
518:Oviposition
456:Caterpillar
417:oviposition
363:iridescence
228:Subspecies
156:Lepidoptera
61:Oxfordshire
44:Oxfordshire
2633:Gonepteryx
2622:Categories
2591:Q109585204
2512:GonepRhamn
1074:References
1034:monandrous
957:Physiology
946:monandrous
894:wavelength
881:structural
690:Adult male
629:Life cycle
599:cremastral
578:tubercules
545:Life cycle
524:univoltine
468:) and the
394:Palearctic
321:Palearctic
177:Gonepteryx
136:Arthropoda
2220:1523-1739
2168:1365-3032
2125:0937-7409
2067:0937-7409
1863:1366-638X
1799:0013-8894
1747:0029-8549
1709:Oecologia
1657:0013-886X
1571:1365-2311
1286:0013-8894
1240:1365-2311
1176:0028-1042
1117:cite book
1109:641575222
1047:G. rhamni
978:Olfaction
972:G. rhamni
835:G. rhamni
820:braconids
814:Parasites
800:G. rhamni
794:Predators
772:Migration
538:parasites
533:predators
528:G. rhamni
429:evergreen
425:hibernate
377:in 1758.
347:buckthorn
333:brimstone
329:its genus
313:butterfly
184:Species:
122:Kingdom:
116:Eukaryota
2585:Wikidata
2509:MaBENA:
2487:LepIndex
2479:10771686
2346:BioLib:
2317:Wikidata
2228:19637406
2133:22444773
2075:21156264
1929:18089132
1763:19024014
1755:24817198
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1579:86578856
1445:17669418
1248:83866795
1192:14494428
1184:25280559
1057:See also
1030:diapause
1011:Diapause
1005:foraging
994:and the
984:antennal
934:latitude
890:pigments
594:Pupation
413:wetlands
400:to east
369:Taxonomy
317:Pieridae
288:Linnaeus
275:Synonyms
216:Linnaeus
166:Pieridae
162:Family:
132:Phylum:
126:Animalia
112:Domain:
89:IUCN 3.1
2605:4535267
2453:1920712
2323:Q206474
2200:Bibcode
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2021:3545080
2001:Bibcode
1975:4199869
1955:Bibcode
1901:Bibcode
1871:1653379
1717:Bibcode
1551:Bibcode
1515:2404579
1495:Bibcode
1417:Bibcode
1370:3546246
1350:Bibcode
1220:Bibcode
1156:Bibcode
996:terpene
885:pigment
859:cryptic
789:Enemies
613:sulphur
574:instars
408:stage.
382:sulphur
311:, is a
172:Genus:
152:Order:
146:Insecta
142:Class:
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962:Vision
940:Mating
583:midrib
437:nectar
406:larval
398:Europe
2538:78609
2499:LoB:
2474:IRMNG
2466:52771
2414:91102
2409:EUNIS
2375:6L326
2362:73918
2349:51362
2129:S2CID
2071:S2CID
2017:JSTOR
1993:Oikos
1971:S2CID
1889:(PDF)
1867:S2CID
1759:S2CID
1705:(PDF)
1575:S2CID
1511:JSTOR
1405:(PDF)
1366:JSTOR
1342:Oikos
1244:S2CID
1188:S2CID
864:rigid
808:wasps
804:birds
606:Adult
552:imago
480:Adult
433:holly
42:Pit,
2600:GBIF
2533:NCBI
2502:6484
2448:GBIF
2396:EPPO
2357:BOLD
2250:ISBN
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1180:PMID
1172:ISSN
1123:link
1105:OCLC
1095:ISBN
883:and
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806:and
736:♂ △
678:Pupa
589:Pupa
497:and
402:Asia
292:1758
220:1758
2551:699
2520:NBN
2383:EoL
2370:CoL
2208:doi
2156:doi
2113:doi
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2009:doi
1963:doi
1951:205
1917:hdl
1909:doi
1851:doi
1733:hdl
1725:doi
1713:175
1649:101
1559:doi
1503:doi
1433:hdl
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