See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/223001066 Kleptoparasitism in birds ARTICLE in ANIMAL BEHAVIOUR · MAY 1979 Impact Factor: 3.14 · DOI: 10.1016/0003-3472(79)90185-4 CITATIONS READS 292 625 2 AUTHORS, INCLUDING: H. Jane Brockmann University of Florida 75 PUBLICATIONS 2,425 CITATIONS SEE PROFILE All in-text references underlined in blue are linked to publications on ResearchGate, Available from: H. Jane Brockmann letting you access and read them immediately. Retrieved on: 30 January 2016 Anim. Behav., 1979,27,487-514 KLEPTOPARASITISM IN BIRDS BY H. JANE BROCKMANN* & C. J. BARNARD? Animal Behaviour Research Group, Department of Zoology, University of Oxford Abstract. Kleptoparasitism refers to the interspecific stealing of already procured food, but this paper shows that, intraspecific food-stealing is effectively the same behaviour. A comprehensive review of the literature shows that certain orders of birds contain a disproportionate number of kleptoparasitic species. Birds in these orders occupy a limited range of ecological niches and are most commonly either predatory or dietary opportunists. Kleptoparasitism is particularly associated with certain ecological conditions, such as the availability of hosts feeding on large, visible food items and periods of food shortage. Birds show a wide range of socially parasitic feeding interactions of which kleptoparasitism is one extreme. The parasitic pattern of food-stealing is likely to involve frequency-dependent selection and may be an example of an evolutionarily stable strategy. The term ‘clepto-parasitism’ or kleptoparasitism hypotheses to account for the distribution of was introduced by Rothschild & Clay (1952) kleptoparasitism among birds or to demonstrate to describe the stealing of already procured food its relationship to other forms of intra- and inter- by individuals of one species from individuals specific feeding. In this paper we review the of another. The same behaviour is also referred occurrence of kleptoparasitism among birds to as ‘piracy’ by many authors (Meinertzhagen and the ecological conditions under which the 1959, 1964; Ashmole 1971; Nakamura 1972; behaviour emerges. We discuss food stealing Hatch 1975; Andersson 1976; Kgllander 1977) as one of a number of feeding interactions. We as well as ‘food parasitism’ (Hopkins & Wiley also discuss the selective pressures acting to 1972), ‘pilfering’ (Rand 1954) and ‘robbery’ maintain kleptoparasitism as a feeding pattern (Hulsman 1976). Although kleptoparasitism is in a population. a term that is generally reserved for interspecific stealing of food, the other terms are also used Taxonomic Distribution of lrueptoparasitism for intraspecific food theft, emphasizing the in Birds close relationship between intra- and interspecific We have reviewed the ornithological literature feeding patterns. Kleptoparasitic behaviour is of the past 40 years, searching for descriptions known to occur in other animal groups (Wilson of behaviour that could be called klepto- 1971), but it appears to be particularly wide- parasitic. Although undoubtedly incomplete, spread among birds. Species such as frigate- this survey is representative of the species which birds (Fregata spp.) (Bent 1923; Nelson 1975), either kleptoparasitize or act as hosts. The skuas (Stercorarius spp.) (Meinertzhagen 1959; survey (Appendices A, B, C) reveals that birds Grant 1971; Ramsey 1973; Andersson 1976; of some orders are more likely to develop Furness 1977; Arnason & Grant 1978) and some kleptoparasitic habits than those of other orders gulls (Larus spp.) (Schmidt 1954; Bergman (Table I). For example, only one duck, the 1960; Ingolfsson 1969; Hatch 1970; Fuchs 1977; American wigeon (Anas americana) is a Kgllander 1977) often procure a significant regular kleptoparasite, although ducks are proportion of their diet through kleptoparasitism. frequently found in mixed-species flocks and are Occasional acts of piracy have been observed the hosts of other kleptoparasites (Table I). in a wide variety of other bird species (see Among such seed- or fruit-eating orders as Appendix A). Galliformes? Columbiformes, and Psittaciformes, Most of the recorded instances of klepto- kleptoparasltlsm has never been observed. parasitism include only descriptions of the Among the Falconiformes and Charadriiformes behaviour in a particular species, with little (particularly Stercorariidae and Laridae), many attempt to provide an explanation. Only Rand species have been observed kleptoparasitizing (1954) has provided any broadly applicable other birds. Although the vast majority of bird species are in the order Passeriformes, the *Present address: Dept. of Zoology, University of Florida, Gainesville, Florida 32611. incidence of kleptoparasitism between passerines tPresent address: Dept. of Zoology, University of and other birds is infrequent and sporadic. Nottingham. However, passerines are known to steal from 487 488 ANIMAL BEHAVIOUR, 27, 2 Table I. The Oeewrence of Rleptoparasltlsmbetween Birds No. klepto- No. species No. species parasitic acting as kleptopara- No. species speciesin hosts in sitizing Taxon in taxon taxoll taxon thesehosts Sphenisciformes Spheniscidae 18 3 6 0 0 Rheiformes 2 0 0 4 0 0 3 0 0 0 Tinamiformes 45 0 0 0 Gaviiformes Gaviidae 4 0 3 4 Podicipediformes Podicipedidae 19 0 3 5 Procellariiformes 99 12 Diomedeidae Procellariidae :: 1; Hydrobatidae 20 Pelecanoididae 5 :, 58 10 23 Phaethontidae 3 Pekcanidae : : Sulidae ; 4 Phalacroccracidae 29 f Anhingidae 4 i 0 Fregatnlae 5 1 1 Ci~diii~es 123 20 67 14 Cochleariidae 1 0 Balaenicipitidae 1 Scopidae 1 8 Ciconiidae Threskiomithidae :: : Phoenicopteridae 5 0 160 17 47 Anatidae 15: 1’: 4’: Falccniformes 298 31 26 60 Cathartidae 7 1 Saggittariidae : :. Accipitridae 22; 2: 19 Pandionidae ‘! Falccnidae 6: : i 16 Galliformes 270 0 0 0 Gruifoqnes 201 16 ti : ;-;;p 0 Rallidae 1:: 3 s Otididae 25 (other families) 15 ii i chJi&adlgemes 32 47 13; Rostratulidae 0” 0” 0 Haematopodidae 0 1 3 BROCKMANN & BARNARD: KLEPTOPARASITISM IN BIRDS 489 Table I continued No. klepto- No. species No. species parasitic acting as w”nv& N; y&a speciesin hosts in Taxon taxon taxon these hosts Charadriidae 1 Scolopacidae z Recurvirostridae 00 0 Phalaropodidae Dromadidae x 8 Burhinidae Glareolidae 00 00 Thinocoridae Chionididae i 8 Stercorariidae Laridae 2: 9: ;gihzpidae 0 1 2: Columbiformes 313 Psittaciformes 327 Musophagiformes 23 CUCUlifOlTII~ 132 Strigiformes 134 3 5 Tytonidae Strigidae 12; :. : Caprimulgiformes 98 0 0 Apodiformes 385 0 0 Coliiformes 6 0 0 0 Trogoniformes 36 0 0 0 Coraciiformes 205 Alcedinidae 91 Todidae 5 Momotidae Meropidae 2: Coraciidae 16 Leptosomatidae 1 Upupidae Phoeniculidae ; Bucerotidae 50 Piciformes 397 0 0 0 Passerifqrmes* 5266 23 23 42 ye;;y 1;: : : A- Paridae 62 Sittidae 22 0” : :. Cinclidae 1 g@;&$idae 6: 8 : : Turdidae 3:; : 7 1; Motacillidae 1 Laniidae t: : Stumidae 1 : Icteridae !I Ploceidae : s Fringillidae 2 : 2 *Only the families in which kleptoparasitism occursare included. 490 ANIMAL BEHAVIOUR, 27, 2 insects and spiders’ webs more frequently than A) and from mammals (Appendix B). This are other species (Appendix C). The frequency behaviour usually involves chasing or directly of kleptoparasitism within an order does not snatching the prey from the other predator. appear to be simply a function of the number of species. Rather there appear to be common 2. Association Based on Mobbing features between orders which make them more When birds mob a predator, it may be carry- prone to develop kleptoparasitism. It is easy to ing or feeding on a prey item. If the mobbing see, for example, that kleptoparasitism is more is successful and the predator driven away, food common among predatory species such as may be left behind which the birds can then eat. raptors and gulls than among non-predatory Such appears to be the probable origin of species. What then are the patterns to the in- magpies (Pica pica) stealing prey from a golden cidence of kleptoparasitism? eagle (Aquila chrysaetos) (Dixon 1933) and of crows (Corvus spp.) stealing from marsh hawks Interspecific Associations that May Lead to (Circus cyaneus), falcons (Falco spp.) and kites KIeptoparasitism (Milvus migrans and Haliastur sphenurus) (Bent Association between two species is obviously 1937; Tinning & Tinning 1970; Balfour 1973; a prerequisite for the development of klepto- Goodwin 1976). parasitic behaviour. Interspecific associations occur: (1) where predators pursue prey, (2) 3. Association Based on Mixed-Species Flocks where the members of one species mob a Kleptoparasitism is common among species predatory animal, (3) where individuals forage which occur in mixed flocks (Rand 1954). in mixed-species flocks, (4) where individuals Birds in these groups aggregate because of a nest in mixed-species aggregations such as common attraction to a rich food source or seabird colonies, (5) where animals join others because they share the same habitat (Rand 1954; in specialized feeding associations such as where Friedmann 1967). In winter, migrant sandpipers, one species acts as a ‘beater’ for another (Rand herons and ducks feed together along shorelines 1954) and (6) where members of one species and on mudflats (Williams 1953; Meyerriecks feed on the products, scraps or parasites of & Nellis 1967; Recher & Recher 1969; Rees another (Rand 1954). 1969; Goss-Custard 1970; Haverschmidt 1970; Siegfried & Batt 1972; Reynolds 1974a; Dawson 1. Association Based on Predation 1975; Raffaele