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Mud-puddling, or simply puddling, is a behaviour most conspicuous in
butterflies
Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises t ...
, but occurs in other animals as well, mainly insects; they seek out
nutrients
A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excr ...
in 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
Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
, dung or
carrion
Carrion () is the decaying flesh of dead animals, including human flesh.
Overview
Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
.
[ (1996): Mating systems and sexual division of foraging effort affect puddling behaviour by butterflies. ''Ecological Entomology'' 21(2): 193-197]
PDF fulltext
/ref> From the fluids they obtain salt
Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quanti ...
s and amino acids that play various roles in their physiology, ethology
Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objective ...
and ecology.[ (1999): Mud-puddling behavior in tropical butterflies: In search of proteins or minerals? '']Oecologia
''Oecologia'' is an international peer-reviewed English-language journal published by Springer since 1968 (some articles were published in German or French until 1976). The journal publishes original research in a range of topics related to plant ...
'' 119(1): 140–148. (HTML abstract
PDF fulltext
This behaviour also has been seen in some other insects, notably the leafhopper
A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and ...
s, e.g. the potato leafhopper, '' Empoasca fabae''.
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
''Halpe'' is a genus of grass skippers in the family Hesperiidae. It is found in the Indomalayan realm.
Species
*''Halpe albicilla'' de Jong & Treadaway, 1993 Sulawesi.
*''Halpe arcuata'' Evans, 1937 Northeast India, Burma to Malay Peninsula, L ...
''.[ (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
Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in th ...
and tears
Tears are a clear liquid secreted by the lacrimal glands (tear gland) found in the eyes of all land mammals. Tears are made up of water, electrolytes, proteins, lipids, and mucins that form layers on the surface of eyes. The different types of ...
. 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
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
puddling behaviour is more commonly seen in males; for example, '' Speyeria mormonia'' males puddle with a much higher frequency than females. The presence of an assembly of butterflies on the ground acts on ''Battus philenor
''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many di ...
'', for example, as a stimulus to join the presumptive mud-puddling flock.
On soil
In tropical India
India, officially the Republic of India ( Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the ...
this phenomenon is mostly seen in the post-monsoon
A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal osci ...
season. The groups generally include several species, particularly members of the families Papilionidae and Pieridae.
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
A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
during mating as a nuptial gift. This nutrition also enhances the survival rate of the eggs.
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.
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
Papilio glaucus-male puddling.webm, '' Papilio glaucus'' mud-puddling
Other sources of liquid nutrient
Some Orthoptera
Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grass ...
– e.g. the yellow-spined bamboo locust (''Ceracris kiangsu
''Ceracris kiangsu'' is a species of grasshoppers in the subfamily Oedipodinae, sometimes called the yellow-spined bamboo locust. It occurs in Indo-China and southern China, where it may become a locally significant agricultural pest. No subspec ...
'') – are attracted to human urine, specifically to the sodium
Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...
and ammonium
The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternar ...
ions in it. Those Lepidoptera that are attracted to dung (e.g. ''Zeuxidia
''Zeuxidia'' is a genus of very large butterflies in the family Nymphalidae. They are "leaf" butterflies with a cryptic ventral pattern. The genus is Indomalayan ( Burma through Indochina and the Philippines to Sumatra, Java and Borneo.
S ...
'' spp.) or carrion seem to prefer ammonium ions rather than sodium. In rotting, the tissues of fruits release sugars and other organic compounds such as alcohol
Alcohol most commonly refers to:
* Alcohol (chemistry), an organic compound in which a hydroxyl group is bound to a carbon atom
* Alcohol (drug), an intoxicant found in alcoholic drinks
Alcohol may also refer to:
Chemicals
* Ethanol, one of sev ...
s that result from the metabolic processes of decay organisms, used as fuel by butterflies.
In Borneo lowland rain forest, numerous species of butterflies regularly visit decaying fruit to drink. This behavior is mainly opportunistic
Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term ...
, though some are highly attracted to old fruit, notably Satyrinae
The Satyrinae, the satyrines or satyrids, commonly known as the browns, are a subfamily of the Nymphalidae (brush-footed butterflies). They were formerly considered a distinct family, Satyridae. This group contains nearly half of the known diver ...
(e.g. ''Neorina lowii'') and Limenitidinae such as ''Bassarona dunya
''Bassarona dunya'', the great marquis, is a butterfly of the family Nymphalidae. It is found in South-East Asia
Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SE ...
''.
Carrion
Carrion () is the decaying flesh of dead animals, including human flesh.
Overview
Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures ...
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
In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
, e.g. brush-footed butterflies such as ''Cirrochroa
''Cirrochroa'', commonly called yeomen, is a genus of butterflies of the subfamily Heliconiinae in the family Nymphalidae found in southeast Asia. The genus ranges from India to New Guinea.
Species
In alphabetical order:847]"">"''Cirrochroa' ...
emalea'' of the Nymphalinae
The Nymphalinae are a subfamily of brush-footed butterflies (family Nymphalidae). Sometimes, the subfamilies Limenitidinae, and Biblidinae are included here as subordinate tribe(s), while the tribe Melitaeini is occasionally regarded as a distinc ...
or the tawny rajah
''Charaxes bernardus'', the tawny rajah, is a butterfly that belongs to the rajahs and nawabs group, that is, the Charaxinae group of the brush-footed butterflies family. This species can be found in India, China, Indomalaya, and onwards to Indo ...
(''Charaxes bernardus'') of the Charaxinae, as well as gossamer-winged butterflies like '' Curetis tagalica'' of the Curetinae or the common imperial
''Cheritra freja'', the common imperial, is a small butterfly found in India, Myanmar, Malaysia and Sri Lanka that belongs to the gossamer-winged butterflies family (Lycaenidae).
Subspecies
The subspecies of ''Cheritra freja'' are-
* ''Cheri ...
(''Cheritra freja'') of the Theclinae.
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
''Charaxes bernardus'', the tawny rajah, is a butterfly that belongs to the rajahs and nawabs group, that is, the Charaxinae group of the brush-footed butterflies family. This species can be found in India, China, Indomalaya, and onwards to Indo ...
'') or opportunistic (some other '' Charaxes'' and '' Polyura'') tend to have a markedly larger bulk and smaller wings, making them more dashing, maneuvrable flyers than fruit specialists like ''Prothoe franck
''Prothoe franck'', the blue begum, is a butterfly species found in Assam in north-eastern India and throughout a large part of South-East Asia."''Prothoe''_Hübner,_ '' and opportunistic fruit visitors such as ''Charaxes durnfordi">824]"at_Markku_Savela's_''Lepidoptera_and_Some_Other_Life_Forms''
It_bel ...
'' 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
''Limenitis'' is a genus of brush-footed butterflies, commonly called the admirals. The sister butterflies (''Adelpha'') and commander butterflies (''Moduza
''Moduza'' is a genus of south-east Asian (Indomalayan realm) Nymphalidae, brush-foot ...
''),
and some which are attracted to any pungent substance.
Certain
moth
Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of ...
s, mainly of the
subfamily Calpinae
The Calpinae are a subfamily of moths in the family Erebidae described by Jean Baptiste Boisduval in 1840. This subfamily includes many species of moths that have a pointed and barbed proboscis adapted to piercing the skins of fruit to feed on ...
, are somewhat notorious for their blood- and tear-drinking habits. ''
Hemiceratoides hieroglyphica
''Hemiceratoides hieroglyphica'' is a moth from Madagascar. It was discovered in 2006 that it frequents sleeping birds at night, and drinks their tears, using a specialized, harpoon-like proboscis.Hilgartner, R., M. Raoilison, W. Büttiker, D.C. ...
'' of
Madagascar
Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
has been noted to visit and suck tears by inserting their proboscis into the closed eyelids of roosting birds. Similar behaviour has been reported in ''
Azeta melanea
''Azeta'' is a genus of moths in the family Erebidae.
Species
* ''Azeta caudalis'' Felder & Rogenhofer, 1874
* ''Azeta ceramina'' (Hubner, 1821)
* ''Azeta leucoma'' Felder & Rogenhofer, 1874
* '' Azeta melanea'' (Stoll, 1782)
* ''Azeta mimica'' ...
'' in Columbia and ''
Gorgone macarea
Gorgone may refer to:
*''Gorgone'', an album by a jam project featuring Buckethead, Travis Dickerson, and Pinchface
Michael Andrew Hakopian, better known as Pinchface is the drummer of the Deli Creeps, Giant Robot II, and the Cornbugs. He ha ...
'' 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]
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Some species of the
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
''
Calyptra'' are called "vampire moths" as they suck blood from sleeping
vertebrate
Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, including humans. Ophthalmotropy (eye-attraction) and lachryphagy (tear drinking) occur in a number of unrelated moths that visit
mammals. ''
Mecistoptera griseifusa
''Mecistoptera griseifusa'' is a moth of the family Noctuidae first described by George Hampson in 1893. It is found in Sri Lanka
Sri Lanka (, ; si, ශ්රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங் ...
'' 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
spermatophore
A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
s during sexual reproduction.
Tear-drinking is not limited to
moth
Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of ...
s, but has recently also been observed in
cockroach
Cockroaches (or roaches) are a Paraphyly, paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are we ...
es.
This behaviour might thus be far more common than previously thought.
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
''Euthalia aconthea'', the common baron, often called simply baron, is a medium-sized nymphalid butterfly native to Sri Lanka, India and southeast Asia. It flies with stiff wing beats and often glides. The wing is not flapped very far below the ...
'') sipping from a guava fruit
Catagramma pygas-thamyras.webm, ''Catagramma pygas
''Callicore pygas'', the Godart's numberwing or pygas eighty-eight, is a species of butterfly of the family Nymphalidae. It is found in Venezuela, Guyana, Ecuador, Peru, Bolivia, Paraguay and the upper Amazonian region of Brazil.
Description
Its ...
'' drinking from metal fence
References
Insect behavior
Butterflies
Habitats
Ecological restoration
Hydrogeology