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Puddling (biology) : ウィキペディア英語版
Mud-puddling

Mud-puddling, or simply puddling, is behaviour most conspicuous in butterflies, but occurs in other animals as well, mainly insects; they seek out certain moist substances such as rotting plant matter, mud and carrion and they suck up the fluid. Where the conditions are suitable conspicuous insects such as butterflies commonly form aggregations on wet soil, dung or carrion.〔 (1996): Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies. ''Ecological Entomology'' 21(2): 193-197. (PDF fulltext )〕 From the fluids they obtain nutrients such as salts and amino acids that play various roles in their physiology, ethology and ecology.〔 & (1991) Mud puddling by butterflies is not a simple matter Ecological Entomology 16(1):123-127 (PDF fulltext )〕〔 (1999): Mud-puddling behavior in tropical butterflies: In search of proteins or minerals? ''Oecologia'' 119(1): 140–148. (HTML abstract) (PDF fulltext )〕 This behaviour also has been seen in some other insects, notably the leafhoppers, e.g. the potato leafhopper, ''Empoasca fabae''.〔 (1982): Nocturnal occurrences of leafhoppers (Homoptera: Cicadellidae) at soil. ''Journal of the Kansas Entomological Society'' 55(1): 73–74. (HTML abstract )〕
Lepidoptera (butterflies and moths) are diverse in their strategies to gather liquid nutrients. Typically, mud-puddling behavior takes place on wet soil. But even sweat on human skin may be attractive to butterflies such as species of ''Halpe''.〔 (1934): On the sexes of some South American moths attracted to light, human perspiration and damp sand. ''Entomologist'' 102: 769-791.-- Volume 67 pp.81-87〕〔 (2006): Diversity and ecology of carrion- and fruit-feeding butterflies in Bornean rain forest. ''Journal of Tropical Ecology'' 22(1): 25–33. (HTML abstract)〕 More unusual sources include blood and tears. Again, similar behaviour is not limited to the Lepidoptera, and for example, the various species of bees commonly called sweat bees are attracted to various kinds of sweat and tears, including that of humans, and other bee species have been recorded as doing so to various degrees.
In many species puddling behaviour is restricted to males, and the presence of an assembly of butterflies on the ground acts on ''Battus philenor'' for example, as a stimulus to join the presumptive mud-puddling flock.〔
==On soil==

In tropical India this phenomenon is mostly seen in the post-monsoon season. The groups generally include several species, particularly members of the families Papilionidae and Pieridae.〔 (2001): Habitat and altitude preferences of butterflies in Aralam Wildlife Sanctuary, Kerala. ''Tropical Ecology'' 42(2): 277-281. (PDF fulltext )〕
Males seem to benefit from the sodium uptake through mud-puddling behaviour with an increase in reproductive success. The collected sodium and amino acids are often transferred to the female with the spermatophore during mating as a nuptial gift. This nutrition also enhances the survival rate of the eggs.〔 (1987): Puddling in butterflies: sodium affects reproductive success in ''Thymelicus lineola''. ''Physiological Entomology'' 12(4): 461–472.〕〔
* (1996): Sodium: a male nuptial gift to its offspring. ''PNAS'' 93(2): 809–813. (PDF fulltext )〕〔 (2004): The effect of male sodium diet and mating history on female reproduction in the puddling squinting bush brown ''Bicyclus anynana'' (Lepidoptera). ''Behavioral Ecology and Sociobiology'' 56(4): 404–411. (HTML abstract)〕
When puddling, many butterflies and moths pump fluid through the digestive tract and release fluid from their anus. In some, such as the male notodontid ''Gluphisia septentrionis'', this is released in forced anal jets at 3 second intervals. Fluid of up to 600 times the body mass may pass through and males have a much longer ileum (anterior hindgut) than non-puddling females.〔Scott R. Smedley in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of Insects. Academic Press.〕

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