# Puddling (behavior)

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{{Short description|Insect feeding behaviour}}
[[File:Parthenos sylvia-Kadavoor-2016-06-25-001.jpg|thumb|300px|''[Parthenos sylvia](/source/Parthenos_sylvia)'' mud-puddling at the edge of a forest stream]]

'''Puddling''' is a behaviour in which an organism seeks out [nutrients](/source/nutrients) in certain moist substances such as rotting plant matter, [mud](/source/mud), and [carrion](/source/carrion), and sucks up the fluid. Where the conditions are suitable, conspicuous insects such as [butterflies](/source/butterfly) commonly form aggregations on wet [soil](/source/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 [salt](/source/salt_(chemistry))s and [amino acids](/source/amino_acids) that play various roles in their physiology, [ethology](/source/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](/source/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 [leafhopper](/source/leafhopper)s, e.g. the potato leafhopper, ''[Empoasca fabae](/source/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](/source/Lepidoptera) (butterflies and [moths](/source/moths)) are diverse in their strategies to gather liquid nutrients. Typically, mud-puddling behaviour takes place on wet soil. But even [sweat](/source/sweat) on human skin may be attractive to butterflies such as species of ''[Halpe](/source/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](/source/blood) and [tears](/source/Lachryphagy). Again, similar behaviour is not limited to the Lepidoptera; for example, the various species of bees commonly called [sweat bees](/source/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.<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](/source/Lachryphagy) insects, including lepidopterans, [dipterans](/source/dipterans), and bees, drink the tears of other animals.

In many [species](/source/species), puddling behaviour is more commonly seen in males. For example, ''[Speyeria mormonia](/source/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](/source/Battus_philenor)'', for example, as a stimulus to join the presumptive mud-puddling flock.<ref name="Sculley"/>

==On soil==
In tropical [India](/source/India) this phenomenon is mostly seen in the post-[monsoon](/source/monsoon) season. The groups generally include several species, particularly members of the families [Papilionidae](/source/Papilionidae) and [Pieridae](/source/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](/source/spermatophore) during mating as a [nuptial gift](/source/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](/source/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](/source/Proceedings_of_the_National_Academy_of_Sciences)'' '''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](/source/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](/source/Notodontidae) ''[Gluphisia crenata](/source/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](/source/stream_bed)
File:Spot Swordtails mud puddling Drop.jpg|[Spot swordtail](/source/Spot_swordtail) excreting excess water after mud-puddling
File:Papilio glaucus-male puddling.webm|Male ''[Papilio glaucus](/source/Papilio_glaucus)'' mud-puddling
File:Danaus Plexippus Monarch Butterfly.jpg|Male [Monarch butterfly](/source/Monarch_butterfly) mud-puddling
File:Whites puddling.webm|thumbtime=70|Cabbage white (''[Pieris rapae](/source/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](/source/Dryas_iulia)'' butterfly [drinking from the tears](/source/Lachryphagy) of a turtle]]

Some [Orthoptera](/source/Orthoptera) – e.g. the yellow-spined bamboo locust (''[Ceracris kiangsu](/source/Ceracris_kiangsu)'') are attracted to human urine, specifically to the [sodium](/source/sodium) and [ammonium](/source/ammonium) [ion](/source/ion)s 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](/source/Lepidoptera) that are attracted to dung (e.g. ''[Zeuxidia](/source/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](/source/Oecologia)'' '''117'''(4): 536-542. {{doi|10.1007/s004420050690}} (HTML abstract)</ref> In rotting, the tissues of fruits release [sugar](/source/sugar)s and other organic compounds such as [alcohol](/source/Alcohol_(chemistry))s 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.](/source/Biological_Journal_of_the_Linnean_Society)'' '''74''': 1–15. [http://www.urbanwildlands.org/devries/DeVriesWallaBJLS2001.pdf PDF fulltext]</ref>

In [Borneo lowland rain forest](/source/Borneo_lowland_rain_forest), numerous species of butterflies regularly visit decaying fruit to drink. This behavior is mainly [opportunistic](/source/Opportunism), though some are highly attracted to old fruit, notably [Satyrinae](/source/Satyrinae) (e.g. ''Neorina lowii'') and [Limenitidinae](/source/Limenitidinae) such as ''[Bassarona dunya](/source/Bassarona_dunya)''.<ref name="HamerEtAl"/>

[Carrion](/source/Carrion) is usually more intentionally utilized. Carrion-feeders seem to represent a different [feeding guild](/source/feeding_guild) from "classical" mud-puddlers and fruit-feeders. They include diverse [taxa](/source/taxa), e.g. [brush-footed butterflies](/source/brush-footed_butterflies) such as ''[Cirrochroa](/source/Cirrochroa) emalea'' of the [Nymphalinae](/source/Nymphalinae) or the [tawny rajah](/source/tawny_rajah) (''Charaxes bernardus'') of the [Charaxinae](/source/Charaxinae), as well as [gossamer-winged butterflies](/source/gossamer-winged_butterflies) like ''[Curetis](/source/Curetis) tagalica'' of the [Curetinae](/source/Curetinae) or the [common imperial](/source/common_imperial) (''Cheritra freja'') of the [Theclinae](/source/Theclinae).<ref name="HamerEtAl"/>

Carrion-feeding has [evolved](/source/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](/source/Charaxinae), specialist (''[Charaxes bernardus](/source/Charaxes_bernardus)'') or opportunistic (some other ''[Charaxes](/source/Charaxes)'' and ''[Polyura](/source/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](/source/Prothoe_franck)'' and opportunistic fruit visitors such as ''[Charaxes durnfordi](/source/Charaxes_durnfordi)''. Other butterflies like most [Pieridae](/source/Pieridae), [Papilionidae](/source/Papilionidae) and [Morphinae](/source/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](/source/Nymphalidae) show the highest variety of nutrient-gathering strategies among the butterflies; the [Limenitidinae](/source/Limenitidinae) have numerous mud-puddlers that also frequently visit dung but avoid fruits and carrion (namely the genus ''[Limenitis](/source/Limenitis)''),<ref name="HamerEtAl"/> and some which are attracted to any pungent substance.<ref>Several ''[Apatura](/source/Apatura)'' species are famous among [lepidopterologist](/source/lepidopterologist)s for being easily attracted with stinking cheese.</ref>

Certain [moth](/source/moth)s, mainly of the [subfamily](/source/subfamily) [Calpinae](/source/Calpinae), are somewhat notorious for their blood- and tear-drinking habits. ''[Hemiceratoides hieroglyphica](/source/Hemiceratoides_hieroglyphica)'' of [Madagascar](/source/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.](/source/Biology_Letters)'' '''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](/source/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](/source/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](/source/genus) ''[Calyptra](/source/Calyptra_(moth))'' are called "vampire moths" as they suck blood from sleeping [vertebrate](/source/vertebrate)s, including humans. '''Ophthalmotropy''' (eye-attraction) and [lachryphagy](/source/lachryphagy) (tear drinking) occur in a number of unrelated moths that visit [mammal](/source/mammal)s. ''[Lobocraspis griseifusa](/source/Lobocraspis_griseifusa)'' is a notable example.<ref name="Baenziger"/> Stingless bees in the genera ''[Lisotrigona](/source/Lisotrigona)'' and ''[Pariotrigona](/source/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](/source/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 [spermatophore](/source/spermatophore)s 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 [moth](/source/moth)s, but has recently also been observed in [cockroach](/source/cockroach)es.<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](/source/Apatura_ilia)'' f. ''ilia'') on dung
File:Cyclosia papilionaris by Kadavoor.JPG|''[Cyclosia papilionaris](/source/Cyclosia_papilionaris)'' feeding on a bird dropping
File:Common_Baron_DSC_1478.jpg|Common baron (''[Euthalia aconthea](/source/Euthalia_aconthea)'') sipping from a guava fruit
Catagramma pygas-thamyras.webm|''[Catagramma pygas](/source/Catagramma_pygas)'' drinking from metal fence
</gallery>

==References==

{{reflist}}

{{commonscat}}

Category:Insect behavior
Category:Lepidoptera biology
Category:Butterflies
Category:Habitats
Category:Ecological restoration
Category:Hydrogeology

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Adapted from the Wikipedia article [Puddling (behavior)](https://en.wikipedia.org/wiki/Puddling_(behavior)) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Puddling_(behavior)?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.