Kleptoparasitism

There are many lineages of cuckoo bees, all of which lay their eggs in the nest cells of other bees, often within the same family.[3] Bombus bohemicus, for example, parasitises several other species of Bombus, including Bombus terrestris, Bombus lucorum (white-tailed bumblebee), and Bombus cryptarum.[4] The largest monophyletic lineage of kleptoparasitic bees is Nomadinae (a subfamily of Apidae), which comprises several hundred species in 35 genera, all of which are kleptoparasites.[5]

The cuckoo wasps include the family Chrysididae.[6] Many species of chrysidid lay their eggs in the nests of potter and mud dauber wasps. Other families of wasps have "cuckoo" species that parasitise related species, as for example the vespid Polistes sulcifer, which parasitises a related species, Polistes dominula.[7][8] 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). While these insects are usually referred to as kleptoparasites[3] other terms used (not exact synonyms) are inquiline and brood parasite.[9] Another term used is kleptoparasitoid which has been defined as a "parasitoid that increases its reproductive success at the cost of another parasitoid, by preferring hosts that have been parasitized (or sometimes only found) by another female". Kleptoparasitoids may make use of the punctures made by previous parasitoids on their hosts; make use of the trails or traces left by parasitoids to locate hosts; or use hosts that have already been weakened by other parasitoids.[10]

Miltogrammine fly (Craticulina seriata) is a kleptoparasite of sand wasps, depositing its larvae on the food reserved for the larvae of the wasp

Some flies are kleptoparasites, which 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 spider webs where they scavenge on half-eaten stink bugs. Others are associated with robber flies (Asilidae), or Crematogaster ants.[11] Flies in the genus Bengalia (Calliphoridae) steal food and pupae transported by ants and are often found beside their foraging trails.[12] Musca albina (Muscidae) reportedly shows kleptoparasitic behaviour, laying eggs only in dung balls being interred by one out of several co-occurring dung-rolling scarab species.[13]

Scarab dung beetles relocate vast amounts of vertebrate dung to establish their nests. A few species do not transport dung materials, but merely use reserves made by other species, either "roller" or "tunneller". Examples are the genus Cleptocaccobius (small species parasiting balls of roller dung beetles), and the genus Pedaria (whose species nidificate in the nests of large tunneller dung beetles in tropical Africa).

Velia caprai (Ardennes, Belgium)

Many semiaquatic bugs (Heteroptera) are known to engage in kleptoparastism of prey. 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.[14]

Argyrodes flavescens on the web of Argiope pulchella

Kleptoparasitic spiders, which steal or feed on prey captured by other spiders, are known to occur in five families:

• Theridiidae (Argyrodes species)
• Dictynidae (Archaeodictyna ulova)
• Salticidae (species of Portia and Simaetha)
• Symphytognathidae (Curimagua bayano)
• Mysmenidae (Isela okuncana, Kilifia inquilina, and Mysmenopsis species).[15]

This gull is probably attempting to steal food from a sea otter

Western gull (Larus occidentalis) in pursuit of an elegant tern (Thalasseus elegans)

Intraspecific kleptoparasitism often occurs between gulls

Kleptoparasitism is relatively uncommon in birds.[16] However, some non-passerine groups, such as skuas, jaegers and frigatebirds, rely extensively on such behavior to obtain food, and others—including raptors, gulls, terns, coots, and some ducks and shorebirds—will do so opportunistically. Among opportunistic species such as the roseate tern, research has found that parent birds involved in kleptoparasitism are more successful in raising broods than non-kleptoparasitic individuals.[17] Bald eagles have also been seen attacking smaller raptors, such as ospreys, to steal fish away from them.[18] Among passerine birds, there are fewer known examples of kleptoparasitism, though masked shrikes have been recorded stealing food from wheatears,[19] and Eurasian blackbirds have been seen stealing smashed snails from other thrushes.[16]

Flying cormorant has taken a fish from the one in the water

Skuas (including the smaller species known as jaegers in North America) are masters of piracy. Their victims are typically gulls and terns, though other fish-eating species (including auks) are also pursued until they disgorge their catches. The fact that skuas are swift and agile fliers—and that they sometimes gang up on a single victim—aids in their success rate.

During seabird nesting seasons, frigatebirds will soar above seabird colonies, waiting for parent birds to return to their nests 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 have been known to seize tropicbirds by their long tail plumes. Many of the frigatebirds' colloquial names, including man-o'-war bird and pirate of the sea, are a clear reference to this kleptoparasitic behaviour.[20] A study of kleptoparasitism in the magnificent frigatebird suggests that the amount of food obtained by kleptoparasitism may be marginal.[21]

Gulls are known to be perpetrators as well as the victims of kleptoparasitism, with some species frequently exhibiting this behavior, particularly during 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, Heermann's gulls, and laughing gulls are known to steal food from brown pelicans; as the pelicans surface and empty the water from their bills, the gulls lurk nearby and grab escaping food items.[22] Several species of gull also steal food from humans, for example takeaway food at seaside resorts.[23]

Several species of coots and gallinules have been recorded engaging in kleptoparasitism.[24] American coots often feed in the company of other waterfowl species, and occasionally will rob diving ducks—including ring-necked ducks, redheads and canvasback—when they surface with food. Eurasian coots steal from conspecifics, as well as from diving and dabbling ducks, and swans. Allen's gallinules rob both conspecifics and African pygmy geese.

The relationship between spotted hyenas and lions, in which each species steals the other's kills,[25] is a form of kleptoparasitism.[14] Cheetahs are often targets of kleptoparasitism. Bears, coyotes and wolves are very opportunistic and all have this behavior. All hyena species engage in this behavior when they can, and jackals also steal from other carnivores' kills.[26] Sperm whales sometimes steal fish from fishermen's lines, making them the largest of all kleptoparasites.[27]

• 
  A cheetah has killed an impala (and eaten part of it), creating a target for kleptoparasitism
• 
  Three minutes later: this spotted hyena and another are running toward the kill
• 
  Thirty-eight seconds later: The cheetah has fled without resistance. The vultures will also engage in kleptoparasitism: when the hyenas move a piece of the carcass, the vultures will take scraps from the ground.

Human intraspecific kleptoparasitism (humans taking food from other humans) is common in times of famine.

In falconry, humans use trained birds of prey for hunting. Humans use tethered cormorants for fishing.

One researcher saw humans chasing lions from captured prey in Waza National Park, Cameroon in 2006. Interviews with Bororo herdsmen suggest this behaviour, which possibly contributes to declining lion populations in the park, is more widespread than recognised.[28][29]

Various crocodiles will steal food from other animals: the Nile crocodile is known to steal from big cats and hyenas.