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Puddling is a behaviour in which an organism seeks out nutrients in certain moist substances such as rotting plant matter, mud, and carrion, and sucks up the fluid. Where the conditions are suitable, conspicuous insects such as butterfly 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 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 Most conspicuous in butterflies, this behaviour also has been seen in some other animals, primarily insects like the , 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 behaviour 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 Lachryphagy. Again, similar behaviour is not limited to the Lepidoptera; for example, the various species of bees commonly called Halictidae are attracted to various kinds of sweat, including that of humans, and other bee species have been recorded as doing so to various degrees. Lachryphagy insects, including lepidopterans, dipterans, and bees, drink the tears of other animals.
In many species, puddling behaviour is more commonly seen in males. For example, Speyeria mormonia males puddle with a much higher frequency than females.Sculley, Colleen E., and Carol L. Boggs. "Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies." Ecological Entomology 21.2 (1996): 193-197. 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.
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 Notodontidae Gluphisia crenata, 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.
In Borneo lowland rain forest, numerous species of butterflies regularly visit decaying fruit to drink. This behavior is mainly opportunistic, though some are highly attracted to old fruit, notably Satyrinae (e.g. Neorina lowii) and Limenitidinae such as Bassarona dunya.
Carrion is usually more intentionally utilized. Carrion-feeders seem to represent a different feeding guild from "classical" mud-puddlers and fruit-feeders. They include diverse taxa, e.g. brush-footed butterflies such as Cirrochroa emalea of the Nymphalinae or the tawny rajah ( Charaxes bernardus) of the Charaxinae, as well as gossamer-winged butterflies like Curetis tagalica of the Curetinae or the common imperial ( Cheritra freja) of the Theclinae.
Carrion-feeding has evolution independently in several lineages. Specialist carrion-feeders may even have the ability to smell out and home in on rotting meat over hundreds of meters. In the Bornean Charaxinae, specialist ( Charaxes bernardus) or opportunistic (some other Charaxes and Polyura) carrion-feeders tend to have a markedly larger bulk and smaller wings, making them more dashing, maneuverable flyers than fruit specialists like Prothoe franck and opportunistic fruit visitors such as Charaxes durnfordi. Other butterflies like most Pieridae, Papilionidae and Morphinae are rarely if ever seen on carrion, dung and rotting fruit, though they may be avid mud-puddlers in the strict sense. Altogether, the Nymphalidae show the highest variety of nutrient-gathering strategies among the butterflies; the Limenitidinae have numerous mud-puddlers that also frequently visit dung but avoid fruits and carrion (namely the genus Limenitis), and some which are attracted to any pungent substance.Several Apatura species are famous among lepidopterologists for being easily attracted with stinking cheese.
Certain , mainly of the subfamily Calpinae, are somewhat notorious for their blood- and tear-drinking habits. Hemiceratoides hieroglyphica of Madagascar has been noted to visit and suck tears by inserting their proboscis into the closed eyelids of roosting birds.* (2007): Malagasy birds as hosts for eye-frequenting moths. Biology Letters 3(2): 117–120. (HTML abstract) PDF fulltext Similar behaviour has been reported in Azeta melanea in Colombia and Gorgone macarea in Brazil. Other cases of moths drinking human tears have been reported from Thailand. (1992): Remarkable new cases of moths drinking human tears in Thailand (Lepidoptera: Thyatiridae, Sphingidae, Notodontidae). Natural History Bulletin of the Siam Society 40: 101–102. Some species of the genus Calyptra are called "vampire moths" as they suck blood from sleeping , including humans. Ophthalmotropy (eye-attraction) and lachryphagy (tear drinking) occur in a number of unrelated moths that visit . Lobocraspis griseifusa is a notable example. Stingless bees in the genera Lisotrigona and Pariotrigona visit the eyes of mammals and have been known to cause distress to humans. Dryas iulia has also been observed agitating the eyes of caimans and turtles in order to force tear production, which the male butterflies of the species can drink for minerals. The minerals, which can also be obtained from more typical mud-puddling behavior, are used for the butterfly's during sexual reproduction.
Tear-drinking is not limited to , but has recently also been observed in . This behaviour might thus be far more common than previously thought.
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