Kleptoparasitism (originally spelt clepto-parasitism, [1] [2] 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 arthropods, especially bees and wasps, but including some true flies, dung beetles, bugs, and spiders. Cuckoo bees are specialized kleptoparasites which lay their eggs either on the pollen masses made by other bees, or on the insect hosts of parasitoid wasps. 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 skuas, 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.
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. [3] [4] The term denotes a form of parasitism involving theft, from Greek κλέπτω (kléptō, 'steal'). [5] The strategy has been widely studied in birds; in four families, all seabirds, the Fregatidae, Chionididae, Stercoraridae, and Laridae, it occurs in more than a quarter of the species. [6] [4]
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. [3] [6]
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. [7] Bombus bohemicus , for example, parasitises several other species in its genus, including B. terrestris , B. lucorum , and B. cryptarum . [8] 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. [9] [10] The largest monophyletic lineage of kleptoparasitic bees is Nomadinae (a subfamily of Apidae), which comprises several hundred species in 35 genera. [11]
Host genus | Parasite genus |
---|---|
Bombus | Bombus ( Psithyrus ) |
Anthophora | Melecta , Zacosmia |
Amegilla | Thyreus |
Megachile | Coelioxys |
The cuckoo wasps (Chrysididae) 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 Polistes sulcifer , which parasitises a related species, P. dominula . [12] [13] 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). [7] Some of these species are inquilines and brood parasites rather than kleptoparasites. [14]
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. [15]
Some true flies (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 spider webs where they scavenge on half-eaten stink bugs. Others are associated with robber flies (Asilidae), or Crematogaster ants. [16] Flies in the genus Bengalia (Calliphoridae) steal food and pupae transported by ants and are often found beside their foraging trails. [17] Musca albina (Muscidae) reportedly shows kleptoparasitic behaviour, laying eggs only in dung balls being interred by one of several co-occurring dung-rolling scarab species. [18]
Scarab dung beetles 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. [19]
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. [20]
Kleptoparasitic spiders, which steal or feed on prey captured by other spiders, are known to occur in five families:
A few bird species are specialist kleptoparasites, while many others are opportunistic. Skuas (including jaegers) and frigatebirds rely heavily on chasing other seabirds to obtain food. Other species—including raptors, gulls, terns, coots, and some ducks and shorebirds—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. [22] [23] Bald eagles have been seen attacking smaller raptors, such as ospreys, to steal fish from them. [24] Among passerine birds, masked shrikes have been recorded stealing food from wheatears, [25] and Eurasian blackbirds have been recorded stealing smashed snails from other thrushes. [22]
During seabird nesting seasons, frigatebirds 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 sometimes seize tropicbirds 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. [26] However, the amount of food obtained by kleptoparasitism in the magnificent frigatebird may be marginal. [27]
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 brown pelicans surface and empty the water from their bills, they sometimes have their food stolen by Heermann's gulls and laughing gulls, which lurk nearby and grab escaping food items. [28] 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. [29]
The relationship between spotted hyenas and lions, in which each species steals the other's kills, [30] is a form of kleptoparasitism. [20] 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 jackals. [31] Human hunters may commonly take the remains of fresh kills from other carnivores, such as lions and Eurasian lynx. [32] [33] [34] Risso's dolphins 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. [35]
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson characterised parasites as "predators that eat prey in units of less than one". Parasites include single-celled protozoans such as the agents of malaria, sleeping sickness, and amoebic dysentery; animals such as hookworms, lice, mosquitoes, and vampire bats; fungi such as honey fungus and the agents of ringworm; and plants such as mistletoe, dodder, and the broomrapes.
Seabirds 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 environmental problems and feeding niches have resulted in similar adaptations. The first seabirds evolved in the Cretaceous period, and modern seabird families emerged in the Paleogene.
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 tails and long hooked bills. Females have white underbellies and males have a distinctive red gular pouch, which they inflate during the breeding season to attract females. Their wings are long and pointed and can span up to 2.3 metres (7.5 ft), the largest wing area to body weight ratio of any bird.
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 distantly related to auks, and even more distantly to waders. Until the 21st century, most gulls were placed in the genus Larus, but that arrangement is now considered polyphyletic, leading to the resurrection of several genera. An older name for gulls is mews, which is cognate with German Möwe, Danish måge, Swedish mås, Dutch meeuw, Norwegian måke/måse, and French mouette, and can still be found in certain regional dialects.
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as symbionts, and information to serve as a form of communication. Trophallaxis is used by some birds, gray wolves, vampire bats, and is most highly developed in eusocial insects such as ants, wasps, bees, and termites.
In evolutionary ecology, a parasitoid is an organism that lives in close association with its host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionary strategies within parasitism, distinguished by the fatal prognosis for the host, which makes the strategy close to predation.
The fork-tailed drongo, also called the common drongo or African drongo, is a small bird found from the Sahel to South Africa that lives in wooded habitats, particularly woodlands and savannas. It is part of the family Dicruridae and has four recognized subspecies, D. a adsimilis, D. a. apivorus, D. a. fugax and D. a. jubaensis. Like other drongos, the fork-tailed is mostly insectivorous; its diet mainly consists of butterflies, termites, and grasshoppers.
Brood parasitism is a subclass of parasitism and phenomenon and behavioural pattern of animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its own, usually using egg mimicry, with eggs that resemble the host's. The strategy involves a form of aggressive mimicry called Kirbyan mimicry.
The great frigatebird is a large seabird in the frigatebird family. There are major nesting populations in the tropical Pacific Ocean, such as Hawaii and the Galápagos Islands; in the Indian Ocean, colonies can be found in the Seychelles and Mauritius, and there is a tiny population in the South Atlantic, mostly on and around St. Helena and Boatswain Bird Island.
The lava gull, also known as the dusky gull, is a medium-sized gull and a member of the "hooded gull" group. It is most closely related to the Laughing gull and Franklin's gull and is the rarest gull in the world. It is endemic to the Galapagos Islands.
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 the apid subfamily Nomadinae, but is sometimes used in Europe to mean bumblebees (Bombus) in the subgenus Psithyrus. Females of cuckoo bees are easy to recognize in almost all cases, as they lack pollen-collecting structures and do not construct their own nests. They often have reduced body hair, abnormally thick and/or heavily sculptured exoskeleton, and saber-like mandibles, although this is not universally true; other less visible changes are also common.
An obligate parasite or holoparasite is a parasitic organism that cannot complete its life-cycle without exploiting a suitable host. If an obligate parasite cannot obtain a host it will fail to reproduce. This is opposed to a facultative parasite, which can act as a parasite but does not rely on its host to continue its life-cycle. Obligate parasites have evolved a variety of parasitic strategies to exploit their hosts. Holoparasites and some hemiparasites are obligate.
With over 850 species, the genus Nomada is one of the largest genera in the family Apidae, and the largest genus of cuckoo bees. Cuckoo bees are so named because they enter the nests of a host and lay eggs there, stealing resources that the host has already collected. The name "Nomada" is derived from the Greek word nomas, meaning "roaming" or "wandering."
Commonly known as cuckoo wasps or emerald wasps, the hymenopteran family Chrysididae is a very large cosmopolitan group of parasitoid or kleptoparasitic wasps, often highly sculptured, with brilliant metallic colors created by structural coloration. They are most diverse in desert regions of the world, as they are typically associated with solitary bee and wasp species, which are also most diverse in such areas. Their brood parasitic lifestyle has led to the evolution of fascinating adaptations, including chemical mimicry of host odors by some species.
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 freeloader flies or jackal flies. However, because of the conditions under which many species breed out, they also are known as filth flies.
Insect ecology is the interaction of insects, individually or as a community, with the surrounding environment or ecosystem.
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. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.
The magnificent frigatebird, frigate petrel or man o' war is a seabird of the frigatebird family Fregatidae. With a length of 89–114 centimetres and wingspan of 2.17–2.44 m it is the largest species of frigatebird. It occurs over tropical and subtropical waters off America, between northern Mexico and Perú on the Pacific coast and between Florida and southern Brazil along the Atlantic coast. There are also populations on the Galápagos Islands in the Pacific and the Cape Verde islands in the Atlantic. It is one of the fastest birds in the world, flying at speeds of up to 95 miles per hour.
A facultative parasite is an organism that may resort to parasitic activity, but does not absolutely rely on any host for completion of its life cycle.
Ammophila sabulosa, the red-banded sand wasp, is a species of the subfamily Ammophilinae of the solitary hunting wasp family Sphecidae, also called digger wasps. Found across Eurasia, the parasitoid wasp is notable for the mass provisioning behaviour of the females, hunting caterpillars mainly on sunny days, paralysing them with a sting, and burying them in a burrow with a single egg. The species is also remarkable for the extent to which females parasitise their own species, either stealing prey from nests of other females to provision their own nests, or in brood parasitism, removing the other female's egg and laying one of her own instead.