{{Short description|Insect feeding behaviour}} [[File:Parthenos sylvia-Kadavoor-2016-06-25-001.jpg|thumb|300px|''Parthenos sylvia'' mud-puddling at the edge of a forest stream]]
'''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 butterflies commonly form aggregations on wet soil, dung, or carrion.<ref name="Sculley">Sculley, C.E. & Boggs, C.L. (1996): Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies. ''Ecological Entomology'' '''21'''(2): 193-197. [http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/1996%20Sculley%20&%20Boggs%20puddling.pdf PDF fulltext] {{Webarchive|url=https://web.archive.org/web/20121018184543/http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/1996%20Sculley%20%26%20Boggs%20puddling.pdf |date=2012-10-18 }}</ref> From the fluids they obtain salts and amino acids that play various roles in their physiology, ethology, and ecology.<ref>Boggs, C.L. & L.A. Jackson (1991) Mud puddling by butterflies is not a simple matter Ecological Entomology 16(1):123-127 {{doi|10.1111/j.1365-2311.1991.tb00199.x}} [http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/1991%20Boggs%20&%20Jackson%20puddling.pdf PDF fulltext] {{Webarchive|url=https://web.archive.org/web/20121021185431/http://www.stanford.edu/group/CCB/Pubs/Boggs_pdfs/1991%20Boggs%20%26%20Jackson%20puddling.pdf |date=2012-10-21 }}</ref><ref name="Beck1999">Beck, J.; Mühlenberg, E. & Fiedler, K. (1999): Mud-puddling behavior in tropical butterflies: In search of proteins or minerals? ''Oecologia'' '''119'''(1): 140–148. {{doi|10.1007/s004420050770}} (HTML abstract) [http://www.biogeography.unibas.ch/Mitarbeiter/Jan_Beck/JBeck_NLU/abstr_publ/Beck%20etal%20(1999)%20Mud-puddling%20Oecologia.pdf PDF fulltext] {{webarchive|url=https://web.archive.org/web/20110707004200/http://www.biogeography.unibas.ch/Mitarbeiter/Jan_Beck/JBeck_NLU/abstr_publ/Beck%20etal%20%281999%29%20Mud-puddling%20Oecologia.pdf |date=2011-07-07 }}</ref> Most conspicuous in butterflies, this behaviour also has been seen in some other animals, primarily insects like the leafhoppers, e.g. the potato leafhopper, ''Empoasca fabae''.<ref>Adler, P.H. (1982): Nocturnal occurrences of leafhoppers (Homoptera: Cicadellidae) at soil. ''Journal of the Kansas Entomological Society'' '''55'''(1): 73–74. [http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=238722&q=&uid=791271398&setcookie=yes HTML abstract] {{webarchive|url=https://web.archive.org/web/20110520045033/http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=238722&q=&uid=791271398&setcookie=yes |date=2011-05-20 }}</ref>
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''.<ref>Collenette, C.L. (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</ref><ref name="HamerEtAl">Hamer, K.C.; Hill, J.K.; Benedick, S.; Mustaffa, N.; Chey, V.K. & Maryati, M. (2006): Diversity and ecology of carrion- and fruit-feeding butterflies in Bornean rain forest. ''Journal of Tropical Ecology'' '''22'''(1): 25–33. {{doi|10.1017/S0266467405002750}} (HTML abstract)</ref> More unusual sources include blood and tears. Again, similar behaviour is not limited to the Lepidoptera; for example, the various species of bees commonly called sweat bees are attracted to various kinds of sweat, including that of humans, and other bee species have been recorded as doing so to various degrees.<ref name="Hogue1993">{{cite book|author=Charles Leonard Hogue|title=Latin American Insects and Entomology|url=https://books.google.com/books?id=3CTf8bnlndwC&pg=PA457|year=1993|publisher=University of California Press|isbn=978-0-520-07849-9|pages=457–}}</ref><ref name="Morris2004">{{cite book|author=Brian Morris|title=Insects and Human Life|url=https://books.google.com/books?id=ok-3NLX_8GQC&pg=PA280|year=2004|publisher=Berg|isbn=978-1-84520-075-6|pages=280–}}</ref> Lachryphagous 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.<ref name="Sculley"/> 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.<ref name="Sculley"/>
==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.<ref>Sreekumar, P.G. & Balakrishnan, M. (2001): Habitat and altitude preferences of butterflies in Aralam Wildlife Sanctuary, Kerala. ''Tropical Ecology'' '''42'''(2): 277-281. {{usurped|1=[https://web.archive.org/web/20110717104324/http://www.tropecol.com/pdf/open/PDF_42_2/42213.pdf PDF fulltext]}}</ref>
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.<ref>Pivnik, K. & McNeil, J.N. (1987): Puddling in butterflies: sodium affects reproductive success in ''Thymelicus lineola''. ''Physiological Entomology'' '''12'''(4): 461–472.</ref><ref>* Medley S.R. & Eisner, T. (1996): Sodium: a male nuptial gift to its offspring. ''PNAS'' '''93'''(2): 809–813. [http://www.pnas.org/cgi/reprint/93/2/809.pdf PDF fulltext]</ref><ref>Molleman, F.; Zwaan, B.J. & Brakefield, P.M. (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. {{doi|10.1007/s00265-004-0789-2}} (HTML abstract)</ref>
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 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.<ref>Scott R. Smedley in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of Insects. Academic Press.</ref>
<gallery mode="nolines" class="center" heights="160" widths="160"> File:Butterflies mud puddling at Aralam Wildlife Sanctuary, Kerala, India (13).jpg|Aggregation of butterflies mud puddling File:Mud-puddling-Aralam-2016-10-29-001.jpg|Collective of different butterfly species mud-puddling on a damp stream bed File:Spot Swordtails mud puddling Drop.jpg|Spot swordtail excreting excess water after mud-puddling File:Papilio glaucus-male puddling.webm|Male ''Papilio glaucus'' mud-puddling File:Danaus Plexippus Monarch Butterfly.jpg|Male Monarch butterfly mud-puddling File:Whites puddling.webm|thumbtime=70|Cabbage white (''Pieris rapae'') butterflies mud-puddling </gallery>
==Other sources of liquid nutrient==
[[File:A butterfly feeding on the tears of a turtle in Ecuador (cropped to butterfly).jpg|thumb|right|A ''Dryas iulia'' butterfly drinking from the tears of a turtle]]
Some Orthoptera – e.g. the yellow-spined bamboo locust (''Ceracris kiangsu'') are attracted to human urine, specifically to the sodium and ammonium ions in it.<ref>Shen, Ke; Wang, Hao-Jie; Shao, Lin; Xiao, Kai; Shu, Jin-Ping; Xu, Tian-Sen & Li, Guo-Qing (2009): Mud-puddling in the yellow-spined bamboo locust, ''Ceracris kiangsu'' (Oedipodidae: Orthoptera): Does it detect and prefer salts or nitrogenous compounds from human urine? ''Journal of Insect Physiology'' '''55'''(1): 78-84. {{doi|10.1016/j.jinsphys.2008.10.011 }} (HTML abstract)</ref> Those Lepidoptera that are attracted to dung (e.g. ''Zeuxidia'' spp.) or carrion seem to prefer ammonium ions rather than sodium.<ref>* Erhardt, A. & Rusterholz, H.P. (1998): Do Peacock butterflies (''Inachis io'') detect and prefer nectar amino acids and other nitrogenous compounds? ''Oecologia'' '''117'''(4): 536-542. {{doi|10.1007/s004420050690}} (HTML abstract)</ref> In rotting, the tissues of fruits release sugars and other organic compounds such as alcohols that result from the metabolic processes of decay organisms, used as fuel by butterflies.<ref>DeVries, P.J. & Walla, T.R. (2001): Species diversity and community structure in neotropical fruit-feeding butterflies. ''Biol. J. Linn. Soc.'' '''74''': 1–15. [http://www.urbanwildlands.org/devries/DeVriesWallaBJLS2001.pdf PDF fulltext]</ref>
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''.<ref name="HamerEtAl"/>
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.<ref name="HamerEtAl"/>
Carrion-feeding has evolved 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''),<ref name="HamerEtAl"/> and some which are attracted to any pungent substance.<ref>Several ''Apatura'' species are famous among lepidopterologists for being easily attracted with stinking cheese.</ref>
Certain moths, 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.<ref>* Hilgartner, R.; Raoilison, Mamisolo; Büttiker, Willhelm; Lees, David C. & Krenn, Harald W. (2007): Malagasy birds as hosts for eye-frequenting moths. ''Biol. Lett.'' '''3'''(2): 117–120. {{doi|10.1098/rsbl.2006.0581}} (HTML abstract) [http://www.univie.ac.at/evobio/Personal/Krenn/PDF/Hilgartner%20et%20al%20%202007.pdf PDF fulltext]{{dead link|date=January 2018 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> Similar behaviour has been reported in ''Azeta melanea'' in Colombia<ref>{{cite journal|author=Sazima, Ivan|year=2015|title=Save your tears: eye secretions of a Ringed Kingfisher fed upon by an erebid moth|journal=Revista Brasileira de Ornitologia|volume=23|issue=4|pages=392–394|doi=10.1007/BF03544314|doi-access=free|hdl=20.500.12733/1251|hdl-access=free}}</ref> and ''Gorgone macarea'' in Brazil.<ref>{{Cite journal|last=de Lima Moraes|first=Leandro João Carneiro|date=2018-09-17|title=Please, more tears: a case of a moth feeding on antbird tears in central Amazonia|journal=Ecology|volume=100|issue=2|article-number=e02518 |doi=10.1002/ecy.2518|pmid=30222853|s2cid=52290796 |issn=0012-9658}}</ref> Other cases of moths drinking human tears have been reported from Thailand.<ref name="Baenziger">Bänziger, H. (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. {{usurped|1=[https://web.archive.org/web/20110721184646/http://www.aseanbiodiversity.info/Abstract/53001446.pdf PDF fulltext]}}</ref> Some species of the genus ''Calyptra'' are called "vampire moths" as they suck blood from sleeping vertebrates, including humans. '''Ophthalmotropy''' (eye-attraction) and lachryphagy (tear drinking) occur in a number of unrelated moths that visit mammals. ''Lobocraspis griseifusa'' is a notable example.<ref name="Baenziger"/> Stingless bees in the genera ''Lisotrigona'' and ''Pariotrigona'' visit the eyes of mammals and have been known to cause distress to humans.<ref>{{Cite journal|last1=Bänziger|first1=Hans|last2=Boongird|first2=Somnuk|last3=Sukumalanand|first3=Prachaval|last4=Bänziger|first4=Sängdao|date=2009|title=Bees (Hymenoptera: Apidae) That Drink Human Tears|journal=Journal of the Kansas Entomological Society|volume=82|issue=2|pages=135–150|issn=0022-8567|jstor=20621944|doi=10.2317/JKES0811.17.1|s2cid=84853663}}</ref> ''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 spermatophores during sexual reproduction.<ref>{{Cite journal|last=de la Rosa|first=Carlos L|date=2014-05-01|title=Additional observations of lachryphagous butterflies and bees|journal=Frontiers in Ecology and the Environment|volume=12|issue=4|pages=210|doi=10.1890/14.wb.006|bibcode=2014FrEE...12..210D }}</ref>
Tear-drinking is not limited to moths, but has recently also been observed in cockroaches.<ref name="van den Burgde Rueda2021">{{cite journal | last1=van den Burg | first1=Matthijs P. | last2=de Rueda | first2=Javier Aznar González | title=Lachryphagy by cockroaches: reptile tears to increase reproductive output? | journal=Neotropical Biodiversity | date=1 January 2021 | volume=7 | issue=1 | pages=276–278 | eissn=2376-6808 | doi=10.1080/23766808.2021.1953892 | pmid=| url=| doi-access=free | bibcode=2021NeBio...7..276V | hdl=10261/249057 | hdl-access=free }}</ref> This behaviour might thus be far more common than previously thought.
<gallery mode="nolines" class="center" heights="160" widths="160"> File:Grand mars changeant.jpg| Blue lesser purple emperor (''Apatura ilia'' f. ''ilia'') on dung File:Cyclosia papilionaris by Kadavoor.JPG|''Cyclosia papilionaris'' feeding on a bird dropping File:Common_Baron_DSC_1478.jpg|Common baron (''Euthalia aconthea'') sipping from a guava fruit Catagramma pygas-thamyras.webm|''Catagramma pygas'' drinking from metal fence </gallery>
==References==
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Category:Insect behavior Category:Lepidoptera biology Category:Butterflies Category:Habitats Category:Ecological restoration Category:Hydrogeology