![Klepto](https://upload.wikimedia.org/wikipedia/commons/3/38/Klepto.JPG)
Kleptoparasitism (etymologically,
parasitism by
theft
Theft is the act of taking another person's property or services without that person's permission or consent with the intent to deprive the rightful owner of it. The word ''theft'' is also used as a synonym or informal shorthand term for som ...
) is a form of
feeding
Eating (also known as consuming) is the ingestion of food, typically to provide a heterotrophic organism with energy and to allow for growth. Animals and other heterotrophs must eat in order to survive — carnivores eat other animals, herbi ...
in which one animal deliberately takes food from another. The strategy is
evolutionarily stable when stealing is less costly than direct feeding, which can mean when food is scarce or when victims are abundant. Many kleptoparasites are
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s, especially bees and wasps, but including some true flies, dung beetles, bugs, and spiders.
Cuckoo bee
The term cuckoo bee is used for a variety of different bee lineages which have evolved the kleptoparasitic behaviour of laying their eggs in the nests of other bees, reminiscent of the behavior of cuckoo birds. The name is perhaps best applied to ...
s are specialized kleptoparasites which lay their eggs either on the pollen masses made by other bees, or on the insect hosts of
parasitoid wasps
Parasitoid wasps are a large group of hymenopteran superfamilies, with all but the wood wasps (Orussoidea) being in the wasp-waisted Apocrita. As parasitoids, they lay their eggs on or in the bodies of other arthropods, sooner or later causi ...
. They are an instance of
Emery's rule
In 1909, the entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.Emery, C. "Über den Ursprung der dulotischen, parasitischen und m ...
, which states that insect social parasites tend to be closely related to their hosts. The behavior occurs, too, in vertebrates including birds such as
skua
The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the long-tailed skua, the Arctic skua, and the pomarine skua are called ...
s, which persistently chase other seabirds until they disgorge their food, and carnivorous mammals such as
spotted hyenas and
lions. Other species opportunistically indulge in kleptoparasitism.
Strategy
Kleptoparasitism is a feeding strategy where one animal deliberately steals food from another. This may be intraspecific, involving stealing from members of the same species, or interspecific, from members of other species.
The term denotes a form of
parasitism involving
theft
Theft is the act of taking another person's property or services without that person's permission or consent with the intent to deprive the rightful owner of it. The word ''theft'' is also used as a synonym or informal shorthand term for som ...
, from Greek κλέπτω (kléptō, 'steal').
The strategy has been widely studied in
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s; in four families, all
seabird
Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s, the
Fregatidae
Frigatebirds are a family of seabirds called Fregatidae which are found across all tropical and subtropical oceans. The five extant species are classified in a single genus, ''Fregata''. All have predominantly black plumage, long, deeply forked ...
,
Chionididae,
Stercoraridae, and
Laridae
Laridae is a family of seabirds in the order Charadriiformes that includes the gulls, terns, skimmers and kittiwakes. It includes around 100 species arranged into 22 genera. They are an adaptable group of mostly aerial birds found worldwide. ...
, it occurs in more than a quarter of the species.
Such a strategy should only be followed if it is
evolutionarily stable, meaning that it offers a
selective advantage to individuals that practise it. Kleptoparasitism costs time and energy which could otherwise be spent directly on feeding, so this cost must be outweighed by the benefit in energy gained from the stolen food. Mathematical modelling suggests that when food is abundant, ordinary feeding is the best strategy; when food abundance falls below a critical level, kleptoparasitism suddenly becomes advantageous, and aggressive interactions become common. Similarly, when potential victims are rare or widely dispersed, the time needed to find them may not be justified by the food that might be stolen from them, resulting in
frequency-dependent selection Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population.
* In positive frequency-dependent selection, the fitness of a phenotyp ...
.
Taxonomic distribution
Arthropods
Bees and wasps
![Cuckoo bee](https://upload.wikimedia.org/wikipedia/commons/5/51/Cuckoo_bee.jpg)
There are many lineages of
cuckoo bee
The term cuckoo bee is used for a variety of different bee lineages which have evolved the kleptoparasitic behaviour of laying their eggs in the nests of other bees, reminiscent of the behavior of cuckoo birds. The name is perhaps best applied to ...
s, all of which lay their eggs in the nest cells of other
bees, often within the same family.
''
Bombus bohemicus
''Bombus bohemicus'', also known as the gypsy's cuckoo bumblebee, is a species of socially parasitic cuckoo bumblebee found in most of Europe with the exception of the southern Iberian Peninsula and Iceland. ''B. bohemicus'' practices inquilini ...
'', for example, parasitises several other species in its genus, including ''
B. terrestris'', ''
B. lucorum'', and ''
B. cryptarum''. These are instances of
Emery's rule
In 1909, the entomologist Carlo Emery noted that social parasites among insects (e.g., kleptoparasites) tend to be parasites of species or genera to which they are closely related.Emery, C. "Über den Ursprung der dulotischen, parasitischen und m ...
, named for the Italian entomologist
Carlo Emery
Carlo Emery (25 October 1848, Naples – 11 May 1925) was an Italian entomologist. He is remembered for Emery's rule, which states that insect social parasites are often closely related to their hosts.
Early in his career Carlo Emery pursue ...
, which asserts that
social parasites among insects, including kleptoparasites, tend to be closely related to their hosts.
The largest
monophyletic lineage of kleptoparasitic bees is
Nomadinae
Nomadinae is a subfamily of bees in the family Apidae. They are known commonly as cuckoo bees.
This subfamily is entirely kleptoparasitic. They occur worldwide, and use many different types of bees as hosts. As parasites, they lack a pollen-car ...
(a subfamily of
Apidae
Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
), which comprises several hundred species in 35 genera.
The
cuckoo wasp
Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group (over 3000 described species) of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic colors ...
s (Chrysididae) lay their eggs in the nests of
potter and
mud dauber
Mud dauber (or "mud wasp" or "dirt dauber") is a name commonly applied to a number of wasps from either the family Sphecidae or Crabronidae which build their nests from mud; this excludes members of the family Vespidae (especially the subfamily ...
wasps. Other families of
wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s have "cuckoo" species that parasitise related species, as for example ''
Polistes sulcifer'', which parasitises a related species, ''
P. dominula''.
Numerous other wasp families have genera or larger lineages of which some or all members are kleptoparasitic (e.g., the genus ''
Ceropales
''Ceropales'' is a genus of kleptoparasitic spider wasps from the sub-family Ceropalinae of the family Pompilidae
Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopoli ...
'' in
Pompilidae
Wasps in the family Pompilidae are commonly called spider wasps, spider-hunting wasps, or pompilid wasps. The family is cosmopolitan, with some 5,000 species in six subfamilies. Nearly all species are solitary (with the exception of some group-ne ...
and the tribe
Nyssonini
The Nyssonini are a group of cleptoparasitic bembicine wasps generally distinguished by the petiolate second submarginal cell of the forewing and rather strongly sculptured head and mesosoma (a common trait in cleptoparasitic wasps). Most specie ...
in
Crabronidae
The Crabronidae are a large paraphyletic group (nominally a family) of wasps, including nearly all of the species formerly comprising the now-defunct superfamily Sphecoidea. It collectively includes well over 200 genera, containing well over 9 ...
).
Some of these species are
inquiline
In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
s and
brood parasites rather than kleptoparasites.
Others are dubbed kleptoparasitoids, namely
parasitoids that select hosts that have been parasitized by another female. Kleptoparasitoids may make use of the punctures made by previous parasitoids on their hosts; may follow the trails or traces left by parasitoids to locate hosts; or use hosts already weakened by other parasitoids.
Flies
Some
true flies
Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
(Diptera) are kleptoparasites; the strategy is especially common in the subfamily
Miltogramminae of the family
Sarcophagidae
Sarcophagidae () are a family of flies commonly known as flesh flies. They differ from most flies in that they are ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on carrion, dung, decaying material, or o ...
. There are also some kleptoparasites in the
families
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
Chloropidae
The Chloropidae are a family of flies commonly known as frit flies or grass flies. About 2000 described species are in over 160 genera distributed worldwide. These are usually very small flies, yellow or black and appearing shiny due to the virtu ...
and
Milichiidae
Milichiidae are a family of flies. Most species are very small and dark. Details of their biology have not yet been properly studied, but they are best known as kleptoparasites of predatory invertebrates, and accordingly are commonly known as fr ...
. Some adult milichiids, for example, visit
spider web
A spider web, spiderweb, spider's web, or cobweb (from the archaic word '' coppe'', meaning "spider") is a structure created by a spider out of proteinaceous spider silk extruded from its spinnerets, generally meant to catch its prey.
Spi ...
s where they scavenge on half-eaten
stink bugs. Others are associated with robber flies (
Asilidae
The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking hypopharynx. The name "robber flies" reflects their notoriously aggressive pre ...
), or ''
Crematogaster
''Crematogaster'' is an ecologically diverse genus of ants found worldwide, which are characterised by a distinctive heart-shaped gaster (abdomen), which gives them one of their common names, the Saint Valentine ant. Members of this genus are ...
''
ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s. Flies in the genus ''
Bengalia
''Bengalia'' is a genus of blow flies in the family Calliphoridae with one authority considering the genus to belong to a separate family Bengaliidae.Lehrer, A.Z., 2003, Bengaliidae n. fam. Une nouvelle famille de Diptera Cyclorrhapha. Entom. Croa ...
'' (
Calliphoridae
The Calliphoridae (commonly known as blow flies, blow-flies, carrion flies, bluebottles, greenbottles, or cluster flies) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing ba ...
) steal food and pupae transported by
ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s and are often found beside their foraging trails. ''
Musca albina'' (
Muscidae
Muscidae are a family of flies found in the superfamily Muscoidea.
Muscidae, some of which are commonly known as house flies or stable flies due to their synanthropy, are worldwide in distribution and contain almost 4,000 described species i ...
) reportedly shows kleptoparasitic behaviour, laying eggs only in dung balls being interred by one of several co-occurring dung-rolling
scarab species.
Dung beetles
Scarab dung beetle
Dung beetles are beetles that feed on feces. Some species of dung beetles can bury dung 250 times their own mass in one night.
Many dung beetles, known as ''rollers'', roll dung into round balls, which are used as a food source or breeding cha ...
s relocate large amounts of vertebrate dung, rolling balls of the material to their nests for their larvae to feed on. Several smaller species of dung beetle do not gather dung themselves but take it from the nests of larger species. For example, species of ''
Onthophagus'' enter dung-balls while ''
Scarabeus'' beetles are making them.
True bugs
![Velia](https://upload.wikimedia.org/wikipedia/commons/0/00/Velia.caprai.jpg)
Many semiaquatic bugs (
Heteroptera) are kleptoparasitic on their own species. In one study, whenever the bug ''
Velia caprai'' (water cricket) took prey heavier than 7.9 g, other bugs of the same species joined it and successfully ate parts of the prey.
Spiders
![Argyrodes flavescens at Nayikayam Thattu](https://upload.wikimedia.org/wikipedia/commons/c/c0/Argyrodes_flavescens_at_Nayikayam_Thattu.jpg)
Kleptoparasitic
spider
Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, which steal or feed on prey captured by other spiders, are known to occur in five families:
*
Theridiidae
Theridiidae, also known as the tangle-web spiders, cobweb spiders and comb-footed spiders, is a large family of araneomorph spiders first described by Carl Jakob Sundevall in 1833. This diverse, globally distributed family includes over 3,000 sp ...
(''
Argyrodes
''Argyrodes'', also called dewdrop spiders, is a genus of comb-footed spiders that was first described by Eugène Louis Simon in 1864. They occur worldwide, and are best known for their kleptoparasitism. They can spin their own webs, but tend to ...
'' species)
*
Dictynidae
Dictynidae is a family of cribellate, hackled band-producing spiders first described by Octavius Pickard-Cambridge in 1871. Most build irregular webs on or near the ground, creating a tangle of silken fibers among several branches or stems of one ...
(''
Archaeodictyna ulova'')
*
Salticidae
Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
(species of ''
Portia'' and ''
Simaetha'')
*
Symphytognathidae (''
Curimagua bayano'')
*
Mysmenidae (''
Isela okuncana'', ''
Kilifia inquilina'', and ''
Mysmenopsis'' species).
Vertebrates
Birds
A few bird species are specialist kleptoparasites, while many others are opportunistic.
Skua
The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the long-tailed skua, the Arctic skua, and the pomarine skua are called ...
s (including jaegers) and
frigatebird
Frigatebirds are a family of seabirds called Fregatidae which are found across all tropical and subtropical oceans. The five extant species are classified in a single genus, ''Fregata''. All have predominantly black plumage, long, deeply forked ...
s rely heavily on chasing other seabirds to obtain food. Other species—including
raptors,
gull
Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century ...
s,
terns
Terns are seabirds in the family Laridae that have a worldwide distribution and are normally found near the sea, rivers, or wetlands. Terns are treated as a subgroup of the family Laridae which includes gulls and skimmers and consists of ...
,
coot
Coots are medium-sized water birds that are members of the rail family, Rallidae. They constitute the genus ''Fulica'', the name being the Latin term for "coot". Coots have predominantly black plumage, and—unlike many rails—they are usually ...
s, and some
duck
Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form ...
s and
shorebird
245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots
Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s—do so opportunistically. Among opportunists such as the
roseate tern, parent birds involved in kleptoparasitism are more successful in raising broods than non-kleptoparasitic individuals.
Bald eagles have been seen attacking smaller raptors, such as
osprey
The osprey (''Pandion haliaetus''), , also called sea hawk, river hawk, and fish hawk, is a diurnal, fish-eating bird of prey with a cosmopolitan range. It is a large raptor reaching more than in length and across the wings. It is brown o ...
s, to steal fish from them. Among
passerine
A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
birds,
masked shrikes have been recorded stealing food from
wheatear
The wheatears are passerine birds of the genus ''Oenanthe''. They were formerly considered to be members of the Thrush (bird), thrush family, Turdidae, but are now more commonly placed in the Old World flycatcher, flycatcher family, Muscicapid ...
s, and
Eurasian blackbird
The common blackbird (''Turdus merula'') is a species of true thrush. It is also called the Eurasian blackbird (especially in North America, to distinguish it from the unrelated New World blackbirds), or simply the blackbird where this does n ...
s have been recorded stealing smashed
snail
A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class G ...
s from other
thrushes
The thrushes are a passerine bird family, Turdidae, with a worldwide distribution. The family was once much larger before biologists reclassified the former subfamily Saxicolinae, which includes the chats and European robins, as Old World flyca ...
.
During
seabird
Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
nesting seasons, frigatebirds soar above
seabird colonies
Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
, waiting for parent birds to return to their
nest
A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materi ...
s with food for their young. As the returning birds approach the colony, the frigatebirds (which are fast and agile) swoop in to pursue them vigorously; they sometimes seize
tropicbird
Tropicbirds are a family, Phaethontidae, of tropical pelagic seabirds. They are the sole living representatives of the order Phaethontiformes. For many years they were considered part of the Pelecaniformes, but genetics indicates they are most cl ...
s by their long tail plumes. Many of the frigatebirds' colloquial names, including ''
man-o'-war bird'' and ''pirate of the sea'', denote this behaviour.
However, the amount of food obtained by kleptoparasitism in the
magnificent frigatebird
The magnificent frigatebird (''Fregata magnificens'') is a seabird of the frigatebird family Fregatidae. With a length of and wingspan of it is the largest species of frigatebird. It occurs over tropical and subtropical waters off America, bet ...
may be marginal.
Gulls are both perpetrators and victims of opportunistic kleptoparasitism, particularly during the breeding season. While the victim is most often another member of the same species, other (principally smaller) gulls and terns can also be targeted. In the
Americas
The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World.
Along with th ...
,
Heermann's gull
Heermann's gull (''Larus heermanni'') is a gull resident in the United States, Mexico and extreme southwestern British Columbia, nearly all nesting on Isla Rasa in the Gulf of California. They are usually found near shores or well out to sea, very ...
s, and
laughing gull
The laughing gull (''Leucophaeus atricilla'') is a medium-sized gull of North and South America. Named for its laugh-like call, it is an opportunistic omnivore and scavenger. It breeds in large colonies mostly along the Atlantic coast of North Am ...
s steal food from
brown pelican
The brown pelican (''Pelecanus occidentalis'') is a bird of the pelican family, Pelecanidae, one of three species found in the Americas and one of two that feed by diving into water. It is found on the Atlantic Coast from New Jersey to the mout ...
s; as the pelicans surface and empty the water from their bills, the gulls lurk nearby and grab escaping food items.
Great black-backed gull
The great black-backed gull (''Larus marinus'') is the largest member of the gull family. Described by the Cornell Lab of Ornithology as "the king of the Atlantic waterfront", it is a very aggressive hunter, pirate, and scavenger. It breeds on t ...
s are skilled kleptoparasites, stealing from other gulls and from raptors. Several species of gull steal food from humans, for example takeaway food at seaside resorts.
File:Western Gull chasing Elegant Tern.jpg, Western gull
The western gull (''Larus occidentalis'') is a large white-headed gull that lives on the west coast of North America. It was previously considered conspecific with the yellow-footed gull (''Larus livens'') of the Gulf of California. The western ...
in pursuit of an elegant tern
The elegant tern (''Thalasseus elegans'') is a tern in the family Laridae. It breeds on the Pacific coasts of the southern United States and Mexico and winters south to Peru, Ecuador and Chile.
This species breeds in very dense colonies on coa ...
File:Kleptoparasitism Great Cormorant.jpg, The flying cormorant has taken a fish from the one in the water.
Mammals
The relationship between
spotted hyenas and
lions, in which each species steals the other's kills, is a form of kleptoparasitism.
Cheetahs are common targets. Bears, coyotes and wolves are very opportunistic and all have this behavior. All
hyena species engage in this behavior when they can, as do
jackal
Jackals are medium-sized canids native to Africa and Eurasia. While the word "jackal" has historically been used for many canines of the subtribe canina, in modern use it most commonly refers to three species: the closely related black-backed ...
s. Human hunters may commonly take the remains of fresh kills from other carnivores, such as lions and
Eurasian lynx
The Eurasian lynx (''Lynx lynx'') is a medium-sized wild cat widely distributed from Northern, Central and Eastern Europe to Central Asia and Siberia, the Tibetan Plateau and the Himalayas. It inhabits temperate and boreal forests up to an eleva ...
.
Risso's dolphin
Risso's dolphin (''Grampus griseus'') is a dolphin, the only species of the genus ''Grampus''. Some of the closest related species to these dolphins include: pilot whales (''Globicephala'' spp.), pygmy killer whales (''Feresa attenuata''), melon ...
s have been observed charging "head-on" at
sperm whales, causing them to open their mouths; it has been suggested that the observed harassment results in some regurgitation, and that the food is then eaten by the Risso's dolphins. The behaviour is rare and may be opportunistic.
Cheetah with impala kill.jpg, A cheetah
The cheetah (''Acinonyx jubatus'') is a large cat native to Africa and central Iran. It is the fastest land animal, estimated to be capable of running at with the fastest reliably recorded speeds being , and as such has evolved specialized ...
has killed an impala (and eaten part of it), creating a target for kleptoparasitism.
Hyenas at stolen impala kill.jpg, A little later, hyenas have driven off the cheetah and are feeding.
See also
*
Evolutionary models of food sharing Evolutionary biologists have developed various theoretical models to explain the evolution of food-sharing behavior—"the unresisted transfer of food from one food-motivated individual to another"— among humans and other animals.
Models of ...
References
External links
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Parasitology