Green-veined white
Green-veined white | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Lepidoptera |
Family: | Pieridae |
Genus: | Pieris |
Species: | P. napi
|
Binomial name | |
Pieris napi |
The green-veined white (Pieris napi) is a butterfly of the family Pieridae.
Appearance and distribution
A
Some authors consider the
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Pieris napi ♂
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Pieris napi ♂ △
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Pieris napi ♀
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Pieris napi ♀ △
Life cycle and food plants
The eggs are laid singly on a wide range of food plants including
Habitat
P. napi is found in damp, grassy places with some shade, forest edges, hedgerows, meadows and wooded river valleys. The later generations widen their habitat use in the search for alternative food plants in drier, but flowery places. In the Mediterranean the insect is also found in scrub around mountain streams or springs and on floodplains with
Flight times
The generations vary with location, elevation and season. In northern Europe there are two or three generations from April to early September. In warmer areas and in some good years there is a fourth generation. In southern Europe there are three or more partially overlapping generations from March to October.
Seasonal variation
In Great Britain, April, May and June specimens have the veins tinged with grey and rather distinct, but are not so strongly marked with black as those belonging to the second flight, which occurs in late July and throughout August. This seasonal variation, as it is called, is also most clearly exhibited on the underside. In the May and June butterfly (plate 13, left side) the veins below are greenish grey, and those of the hindwings are broadly bordered also with this colour. In the bulk of the July and August specimens (plate 13, right side) only the nervures are shaded with greenish grey, and the nervures are only faintly, or not at all, marked with this colour. Now and then a specimen of the first brood may assume the characters properly belonging to the specimens of the second brood; and, on the other hand, a butterfly of the second brood may closely resemble one of the first brood. As a rule, however, the seasonal differences referred to are fairly constant. By rearing this species from the egg it has been ascertained that part (sometimes the smaller) of a brood from eggs laid in June attains the butterfly stage the same year, and the other part remains in the chrysalis until the following spring, the butterflies in each set being of the form proper to the time of emergence.
Other variation
In the typical form -forma typica- the forewings are creamy-white, irrorated with black towards the base. There is an apical blackish blotch, sometimes broken into several terminal spots ; and a black spot between 3 and 4 (in male sometimes absent) In the female there is a black subdorsal posterior spot, and a dorsal confluent mark. The hindwings are creamy-white, the base black-sprinkled and a black costal spot before the apex (in male sometimes absent). The underside of the hindwings and underside apex of the forewings is pale yellow the veins edged with a shading of fine black lines, in the hindwings more broadly.
The ground colour varies from white to cream, sulphur-yellow, chrome yellow and light hues of buff or brown. The spot markings also vary and may be joined or absent. The vein shading varies in colour and in intensity and the shaded bands may be broad or narrow.
Variants, many named, are described by Röber (Europe), Langham (Ireland) and Anon (Britain) [5] [6] [7]
Behaviour
Senses
Recent research has shown that when males mate with a female, they inject methyl salicylate along with their sperm. The smell of this compound repels other males, thus ensuring the first male's paternity of the eggs—a form of chemical mate guarding.[8]
After a female mates, she will display a mate refusal posture that releases methyl salicylate during a subsequent courtship. The release of this
The adult male of this species has a distinctive odour that resembles
Mating system
In the usually
The amount of ejaculate of virgin males during mating is larger than that of non-virgin males. Females therefore must mate more frequently with non-virgin males in order to obtain the necessary amount of male-derived nutrition.
Sexual cooperation and conflict
In P. napi, the nuptial gift is an example of sexual cooperation towards a common interest of both males and females. The existence of nutrients in the ejaculate is beneficial to the females because it increases female fecundity and longevity, and eventually promotes re-mating. The existence of the anti-aphrodisiac, methyl salicylate, is effective in reducing female harassment by other males.[14]
However, the transfer of this ejaculate can cause a conflict over re-mating due to sperm competition. After a female mates, infertile sperm ejaculated by the male will fill the female's sperm storage organ and prevent her from mating. The amount of infertile sperm stored is correlated with the refractory period of a female after mating. Infertile sperm makes up 90% of the sperm count, showing that males manipulate females by preventing them from mating with another male for a certain period of time. Although polyandry benefits females of P. napi by maximizing the amount of transferred nutrients from the male, the infertile sperm storage prolongs female re-mating.[15]
This refractory period makes it harder for females to mate, and females will continue to have difficulty as their age and mating frequency increase. Males who have recently copulated will not transfer as many nutrients to their next mate, but will spend a longer duration of time for each mating. This increases the mating costs for females because they are spending more time copulating and receiving fewer nutrients from the ejaculate. Males take advantage of this because females do not reduce their mating costs by copulating with virgin males.[16] In addition, males will transfer the most methyl salicylate to their first mate to ensure its paternity. However, a female who mates with a virgin male will have the most difficulty re-mating, therefore delaying her from engaging in the preferred polyandry. Males tailor their ejaculate in the sense that the first ejaculate is meant to prolong the refractory period of the female, and every subsequent ejaculate is meant to maximize efficiency in sperm competition.[9]
Monandry and polyandry
Fecundity benefits of polyandry
Fecundity in P. napi varies with the number of eggs produced as well as the lifespan of the female,[17] and is positively influenced by body mass,[18] other genetic factors independent of body size,[19] and with the degree of polyandry.[20] Fecundity increases with the amount of spermatophore material and ejaculate received from males during mating.[18] In particular, mating with recently unmated males can increase overall female reproductive output, though small females are unable to compensate for the negative effects of size on fecundity by mating multiply.[21][22] However, by mating multiply, polyandrous females have overall higher lifetime fecundity, produce more offspring, and have faster-developing offspring compared to monandrous females.[17][20]
Polyandry versus monandry is a genetically-based mating strategy
Despite the benefits of mating multiply, many female P. napi only mate once, irrespective of the number of high quality, more fit, mates available.[19] Monandry is more common in northern populations of P. napi, and sperm competition is correspondingly lower in the north.[18][20] This polymorphic mating system is partially determined by genetic variability within the females genomes.[19] Monandrous and polyandrous females exhibit different heritable reproductive tactics with monandrous females relying on larval derived resources to realize their fecundity and polyandrous females relying on male donations. When genetically polyandrous females are forced into monandry, due to suboptimal mating conditions, they experience reduced life spans.[18] Monandrous and polyandrous females exhibit different life history strategies: at the start of reproduction females that are monandrous produce more eggs than polyandrous females.[17] It is hypothesized that this life history difference is why monandry is more common in the most northern parts of the species’ range,[17] as early investment in reproduction can be more beneficial with shorter mating seasons.
Maintenance of the monandry/polyandry genetic polymorphism
The maintenance of the two mating systems had been hypothesized to be due to the availability of male nutrients, which can vary within the male-biased operational sex ratio. The male biased sex-ratio can lead to strong competition for mates among the males as well as the males are unlikely to encounter more than one female during the mating season. As a result, females often encounter males that have not recently mated that provide large nuptial gifts. Given the size of the nuptial gift provided by males, means that polyandrous females can substantially increase their lifetime fecundity.[20] However, higher mating rates in females comes with a cost, which can explain the presence of monandrous females in primarily polyandrous populations. Polyandrous females have a higher cost of mating characterized by an increase in time spent looking for mates and time spent mating. This results in a decreased time spent looking for food.[17]
Taxonomy
Some authorities consider P. napi to be a
However, the American butterflies cannot successfully reproduce by laying eggs on the invasive weed garlic mustard, Alliaria petiolata, a fact that threatens their survival as garlic mustard out-competes native mustard plants due to having no biological control species present in North America. In Europe, where garlic mustard is native, 76 things consume it.[23]
Similar species
- Pieris bryoniae
- Pieris ergane
- Pieris krueperi – Krueper's small white
- Pieris rapae – small white
- Pieris oleracea
Subspecies
- Pieris napi napi
- Pieris napi adalwinda (Fruhstorfer, 1909) Finland, Sweden
- Pieris napi meridionalis Heyne & Rühl, 1895 Spain, Italy
- Pieris napi segonzaci (le Cerf, 1923) High Atlas
- Pieris napi maura (Verity, 1911) Glacières de Blida, Algeria
- Pieris napi atlantis (Oberthür, 1923) Azrou, Middle Atlas, Morocco
- Pieris napi flavescens (Wagner, 1903) Mödling, Austria
- Pieris napi lusitanica Lep. Portug. Porto: 2, 1929 De Sousa Portugal
For others see Wikispecies.
Synonyms
- Pieris adalwinda Fruhstorfer, 1909[24]
- Pieris arctica Verity, 1911[24]
- Pieris canidiaformis Drenowsky, 1910[24]
- Pieris dubiosa Röber, 1907[24]
- Pieris flavescens Wagner, 1903[24]
- Pieris meridionalis Heyne, 1895[24]
See also
- Dark-veined white
- Mustard white
- List of butterflies of India (Pieridae)
- List of butterflies of Great Britain
- Species problem
References
- S2CID 7164912.
- ISBN 978-0-8247-8390-7.
- ^ Howe, William H. The Butterflies of North America (Garden City, NY: Doubleday, 1975)
- ^ Driesche, F.V.; Blossey, B.; Hoodle, M.; Lyon, S.; Reardon, R., 2010. Biological Control of Invasive Plants in the Eastern United States. USDA Forest Service. Forest Health Technology Enterprise Team. http://wiki.bugwood.org/Archive:BCIPEUS
- Julius Röber, 1909 Pieridae, pp. 39-74, 374, pls. 17-27. In: Seitz, A. (ed.), Die Groß-Schmetterlinge der Erde. 1. Band. Die palaearctischen Tagfalter. – Stuttgart, Fritz Lehmann.[https://www.zobodat.at/pdf/Seitz-Schmetterlinge-Erde_1_1912_Tafeln_en_0001-0190.pdfpdf
- ^ Langham, C. 1922 Some forms of Pieris napi taken in County Fermanagh. Irish Naturalist 31: 42-45 pdf
- ^ Anon British butterfly aberrations
- S2CID 13702247.
- ^ PMID 15315890.
- ISBN 978-1-4000-8234-6
- ^ S2CID 9096451.
- .
- PMID 14561296.
- PMID 10972120.
- PMID 19640869.
- S2CID 20515742.
- ^ S2CID 23160181. Retrieved 13 March 2020.
- ^ . Retrieved 13 March 2020.
- ^ . Retrieved 13 March 2020.
- ^ S2CID 40165781. Retrieved 13 March 2020.
- S2CID 13168534. Retrieved 13 March 2020.
- S2CID 54389039. Retrieved 13 March 2020.
- ^ Becker, R., Gerber E., Hinz H., Katovich E., Panke B., Reardon R., Renz R., Van Riper L., 2013. Biology and Biological Control of Garlic Mustard. The Forest Technology Enterprise Team. https://www.fs.fed.us/foresthealth/technology/pdfs/FS_garlicmustard.pdf
- ^ a b c d e f "Pieris napi (Linnaeus 1758)". Fauna Europaea. Archived from the original on June 4, 2011. Retrieved 2007-04-27.
Further reading
- Asher, Jim et al. The Millennium Atlas of Butterflies of Britain and Ireland Oxford university Press
- Bowden, S. R.; & Riley, Norman Denbigh (1967): The type-material of Pieris napi pseudorapae Verity. Redia 50, pp. [379-380]
- Bowden, S. R. (Aug 68) Pieris napi in Calabria. Entomologist 101, pp. [180-190]
- Bowden, S. R. (Oct 1970) Polymorphism in Pieris: f. sulphurea in Pieris napi marginalis. Entomologist 103, pp. [241-249]
- Bowden, S. R. (1954) Pieris napi L. f. hibernica Schmidt, eine kuenstliche Aberration? Der gegenwaertige Stand der Frage. Mitt. ent. Ges. Basel (nf)4, pp. [9-15, 17-22]
- Bowden, S. R. (1956) Hybrids within the European Pieris napi L. species-group. Proc. Trans. S. Lond. ent. nat. Hist. Soc. 1954–55, pp. [135-159]
- Bowden, S. R. (1961) Pieris napi L. ab. sulphurea Schoeyen Entomologist 94, pp. [221-226]
- Bowden, S. R. (1962) Übertragung von Pieris napi-Genen auf Pieris bryoniae durch wiederholte Ruckkreuzung. Z. Arbgem. Öst. Ent. 14, pp.
- Bowden, S. R. (1966a) Polymorphism in Pieris Entomologist 99, pp. [174-182]
- Bowden, S. R. (1966b) 'Irregular' diapause in Pieris, with a note on Corsican Pieris brassicae L. Proc. Trans. S. Lond. ent. nat. Hist. Soc. 1966, pp. [67-68]
- Bowden, S. R. (1966c) Pieris napi in Corsica. Entomologist 99, pp. 57–68
- Bowden, S. R. (1970a) What is Pieris dubiosa Warren? Ent. Rec. 82, pp.
- Bowden, S. R. (1970b) Pieris napi L.: speciation and subspeciation. Proc. Trans. Br. Entomol. Nat. Hist. Soc.. 3, pp. [63-70]
- Bowden, S. R. (1971). "'Pieris napi' in America: reconnaissance. Proc". Trans. Br. Entomol. Nat. Hist. Soc. 4: 71–77.
- Bowden, S. R. (1972) 'Pieris napi' in America: genetic imbalance in hybrids. Proc. Trans. Br. Entomol. Nat. Hist. Soc.. 4, pp. [103-117]
- Bowden, S. R. (1975a) Some subspecific and infrasubspecific names in Pieris napi L. Ent. Rec. 87, pp. [153-156]
- Bowden, S. R. (1975b) Relation of Pieris melete Menetries to Pieris napi L.: ssp. melete. Proc. Trans. Br. ent. nat. Hist. Soc. 7, pp. [97-102]
- Bowden, S. R. (1979) Subspecific Variation in Butterflies: Adaptation and Dissected Polymorphism in Pieris (Artogeia) (Pieridae). Journal of the Lepidopterists' Society 33(2), pp. [77-111, 40 f
- Bowden, S. R. (): Sexual mosaics in Pieris. Lep. News 12(1-2), pp. [7-13, 1 tbl, 1 f]
- Bowden, S. R. (): Pieris napi L. (Pieridae) and the Superspecies Concept. Journal of the Lepidopterists' Society 26(3), pp. 170–173
- Bowden, S. R. (1985): Taxonomy for a variable butterfly? [Pieris napi]. Ent. Gaz. 36(2), pp. [85-90]
- Carter, David, 1993 Farfalle e falene Fabbri Editori
- Chew, F.S; Watt, W.B (2006). "The green-veined white (Pieris napiL.), its Pierine relatives, and the systematics dilemmas of divergent character sets (Lepidoptera, Pieridae)" (PDF). Biological Journal of the Linnean Society. 88 (3): 413–435. .
- Chinery, Michael, 1987 Guida degli insetti d'Europa Franco Muzzio Editore
- Chinery, Michael, 1989 Farfalle d'Italia e d'Europa De Agostini/Collins
- Chou Io (Ed.) Monographia Rhopalocerum Sinensium, 1-2
- Dyar, 1903 A List of North American Lepidoptera and Key to the Literature of this Order of Insects Bull. U.S. natn. Mus., 52: xix, 723pp
- Edwards (1869). "Descriptions of new species of diurnal Lepidoptera found within the United States". Trans. Am. Entomol. Soc. 2: 369–376. JSTOR 25076222.
- Eitschberger, 1983 Eitschberger, 1984; Systematische Untersuchungen am Pieris napi-bryoniae-Komplex (s.l.) Herbipoliana 1 (1-2): (1) i-xxii, 1–504, (2) 1-601
- Eitschberger (2001). "Eine neue Unterart von Pieris napi (Linnaeus, 1758) vom Polar Ural". Atalanta. 32 (1/2): 85–88.
- Fruhstorfer, 1909 Neue palaearktische Pieriden Int. ent. Zs. 3 (16): 88 (17 July)
- Hensle, 2001 Zur Frage der subspezifischen Zuordnung von Pieris bryoniae lappona Rangnow, 1935 Atalanta 32 (1/2): 89-95
- Hodges, Ronald W. (ed.), 1983 Check List of the Lepidoptera of America North of Mexico
- Korshunov, Y.P. and Gorbunov, P.Y., 1995 The Butterflies (Rhopalocera) of the Asian part of Russia'Pensoft Digital version Archived 2007-08-31 at the Wayback Machinein English
- Lamas Gerardo, 2004 Atlas of Neotropical Lepidoptera; Checklist: Part 4A; Hesperioidea Papilionoidea
- Leraut, Patrice, 1992 Le farfalle nei loro ambienti Ed. A. Vallardi (ecoguide)
- Linnaeus, 1758 Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Symonymis, Locis. Tomis I. 10th Edition Syst. Nat. (Edn 10) 1
- Lorkovic, Zdravko (1968). "Karyologischer Beitrag zur Frage der Fortpflanzungs verhaltnisse Sudeuropäischer Taxone von Pieris napi (L.). (Lep. Pieridae)". Biol. Glasn. 21: 95–136.
- Mazzei Paolo, Reggianti Diego and Pimpinelli Ilaria Moths and Butterflies of Europe
- Pyle, R. M. National Audubon Society: Field Guide to North American Butterflie1981; ISBN 0-394-51914-0
- Scott, J. A. 1986 The butterflies of North America: a natural history and field guide. Stanford University Press, Stanford, California
- Seppänen, E. J, 1970 Suomen suurperhostoukkien ravintokasvit, Animalia Fennica 14
- Tennent, John, 1996 The butterflies of Morocco, Algeria and Tunisia; ISBN 0-906802-05-9
- Tuzov, Bogdanov, Devyatkin, Kaabak, Korolev, Murzin, Samodurov, Tarasov, 1997 Guide to the Butterflies of Russia and adjacent territories; Hesperiidae, Papilionidae, Pieridae, Satyridae; Volume 1
- Verity, 1908; Verity, [1909]; Verity, 1911; Rhopalocera Palaearctica Iconographie et Description des Papillons diurnes de la région paléarctique. Papilionidae et Pieridae Rhopalocera Palaearctica 1: 86+368pp, 2+12+72pls
- Wynter-Blyth, M. A., 1957 Butterflies of the Indian Region; (1982 Reprint)
External links
- Video footage of mud puddling behaviour.
- Pieridae Holarctinae Photos of imagos and la
- www.schmetterling-raupe.de
- Mario Meier - Europäische Schmetterlinge
- www.eurobutterflies.com
- Moths and Butterflies of Europe and North Africa
- Naturkundliches Informationssystem: Pieris napi napi (Linnaeus, 1758)
- Naturkundliches Informationssystem: Pieris napi flavescens F.Wagner, 1903