Kleptoparasitism (originally spelt clepto-parasitism, meaning "parasitism by theft") is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, such as when food is scarce or when victims are abundant. Many kleptoparasites are , especially bees and wasps, but including some true flies, dung beetles, bugs, and spiders. are specialized kleptoparasites which lay their eggs either on the pollen masses made by other bees, or on the insect hosts of Parasitoid wasp. They are an instance of Emery's rule, which states that insect social parasites tend to be closely related to their hosts. The behavior occurs, too, in vertebrates including birds such as , which persistently chase other seabirds until they disgorge their food, and carnivorous mammals such as and . 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, from Greek κλέπτω (kléptō, 'steal').
The strategy has been widely studied in
; in four families, all
, the
Frigatebird,
Sheathbill,
Skua, and
Laridae, it occurs in more than a quarter of the species.
Such a strategy should be followed only if it is evolutionarily stable, meaning that it offers a Adaptation 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.
Taxonomic distribution
Arthropods
Bees and wasps
There are many lineages of
, all of which lay their eggs in the nest cells of other
, often within the same family.
Bombus bohemicus, for example, parasitises several other species in its genus, including
B. terrestris,
Bombus lucorum, and
Bombus cryptarum.
These are instances of Emery's rule, named for the Italian entomologist
Carlo Emery, 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 (a subfamily of
Apidae), which comprises several hundred species in 35 genera.
The (Chrysididae) lay their eggs in the nests of potter wasp and mud dauber wasps. Other families of 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 in Pompilidae and the tribe Nyssonini in Crabronidae). Some of these species are and rather than kleptoparasites.
Others are dubbed kleptoparasitoids, namely 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. Especially the latter is referred to as pirate parasitism.
Flies
Some
fly (Diptera) are kleptoparasites; the strategy is especially common in the subfamily
Miltogramminae of the family
Sarcophagidae. There are also some kleptoparasites in the families
Chloropidae and
Milichiidae. Some adult milichiids, for example, visit
where they scavenge on half-eaten stink bugs. Others are associated with robber flies (
Asilidae), or
Crematogaster .
[Wild, A.L. & Brake, I. 2009. Field observations on Milichia patrizii ant-mugging flies (Diptera: Milichiidae: Milichiinae) in KwaZulu-Natal, South Africa. African Invertebrates 50 (1): 205–212.[1] ] Flies in the genus
Bengalia (
Calliphoridae) steal food and pupae transported by
and are often found beside their foraging trails.
[Sivinski, J., S. Marshall and E. Petersson (1999) Kleptoparasitism and phoresy in the diptera. Florida Entomologist 82 (2) [2] ] Musca albina (
Muscidae) reportedly shows kleptoparasitic behaviour, laying eggs only in dung balls being interred by one of several co-occurring dung-rolling
Scarabaeidae species.
[Marshall, S.A. & Pont, A.C. (2013). The kleptoparasitic habits of Musca albina Wiedemann, 1830 (Diptera: Muscidae). African Invertebrates 54(2): 427–430.]
Dung beetles
Scarabaeidae 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
Scarabaeus beetles are making them.
True bugs
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
Kleptoparasitic
, which steal or feed on prey captured by other spiders, are known to occur in five families:
Vertebrates
Birds
A few bird species are specialist kleptoparasites, while many others are opportunistic.
(including jaegers) and
rely heavily on chasing other seabirds to obtain food. Other species—including raptors,
,
terns,
, and some
and
—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.
have been seen attacking smaller raptors, such as
, to steal fish from them.
Among
passerine birds,
have been recorded stealing food from
,
and Eurasian blackbirds have been recorded stealing smashed
from other thrushes.
During seabird nesting seasons, frigatebirds soar above seabird colony, waiting for parent birds to return to their 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 by their long tail plumes. The name frigatebird, as well as 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 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, as surface and empty the water from their bills, they sometimes have their food stolen by Heermann's gulls and , which lurk nearby and grab escaping food items. Great black-backed gulls 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 ( Larus occidentalis) in pursuit of an elegant tern ( Thalasseus elegans)
File:Fight, lucha. (50206561093).jpg|Black-headed gull ( Chroicocephalus ridibundus) attempting to steal a fish caught by a common tern ( Sterna hirundo)
File:Kleptoparasitism Great Cormorant.jpg|The flying great cormorant ( Phalacrocorax carbo) has taken a fish from the one in the water.
Mammals
The relationship between
and
, 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. Crab-eating macaques have also exhibited kleptoparasitic behaviors. All
hyena species engage in this behavior when they can, as do
.
[Estes, op. cit., 281–295, 339–346] Human hunters may commonly take the remains of fresh kills from other carnivores, such as lions and
Eurasian lynx.
Risso's dolphins have been observed charging "head-on" at
, 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 has killed an impala (and eaten part of it), creating a target for kleptoparasitism.
Hyenas at stolen impala kill.jpg|A little later, spotted hyena have driven off the cheetah and are feeding.
See also
External links