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Charadriiformes (5%) and Gruiformes (4%). On the other hand, low INP rates were reported for altricial birds: 5% Ciconiiformes, 1 Yo among Passeriformes and.
lbis (2001) 143, 133-1 43

An updated list and some comments on the occurrence of intraspecific nest parasitism in birds YORAM YOM-TOV Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel

This paper presents a list, compiled from the literature, of bird species in which intraspecific nest parasitism (INP) occurs. INP was reported in 234 species: one Struthioniformes, two Tinamiformes, two Procellariiformes, six Podicipediiformes, five Ciconiiformes, one Phoenicopteriformes, 74 Anseriformes, one Falconiformes, 32 Galliformes, eight Gruiformes, 19 Charadriiformes, nine Columbiformes, five Cuculiformes, two Apodiformes, one Coraciiformes and 66 Passeriformes. Hence, INP is very common among precocial species, and less so among altricials. Irrespective of this, most INP birds are colonial.

Intraspecific nest parasitism (INP) in birds was considered to be rare compared with interspecific parasitism (Payne 1977), and in 1980 was reported for only 53 species (Yom-Tov 1980a). The growing awareness of the diversity of breeding strategies increased the list considerably, and by 1989 it more than doubled (MacWhirter 1989, Rohwer & Freeman 1989), exceedmg the number of interspecific parasite species (about 80; Payne 1977). A high rate of INP is expected when host nests are available for extended periods of time or in a limited space. Hence, high INP rates are expected in precocial species, because: (1) most lay larger clutches than do similarly sized altricial species (Ar & Yom-Tov 1978); (2) they start incubating mostly after the last or penultimate egg is laid, thus the nest is left without protection for most of the laying period; (3) parasitic eggs and young among precocial birds require a smaller increase in parental care from hosts than in altricial birds (Sorenson 1992). High INP rates are expected when nests are crowded in a small space, i.e. in colonial species or in high nesting density, and when nesting sites are a limiting factor, as they are among non-excavating hole nesters (Yom-Tov 1980a, Anderson 1982, Rohwer & Freeman 1989). The aim of this paper is to provide an up-to-date list of all species for which I found data on INP, and briefly comment on two factors associated with this phenomenon, namely mode of reproduction and coloniality.

METHODS Data on INP were gathered from the ornithological Email: [email protected]

@ 2001 British Ornithologists' Union

literature, including reviews (Yom-Tov 1980a, Rohwer & Freeman 19893, research papers, books on regional avifaunas (Ali & Ripley 1968-1974, Brown et al. 1982, Harrison 1975, Maclean 1993, Urban et al. 1986) and books dealing with a particular group of birds where INP is known to be common (Goodwin 1977, 1982, Johnsgard 1978, 1986, 1988, Summers-Smith 1988). A species was considered to have INP if the above sources described it as such, or if it is stated that more than one female laid in one nest but only one female cared for the young, or if abnormal clutch size was found (larger than twice mean clutch size; Yom-Tov 1980a). These data were complemented with data on nest type (cavity or other), and nesting coloniality (Cramp & Simmons 1977, 1980, Cramp & Perrins 1994, Hoyo et al. 1992, 1994a, 1994b). A species was defined as colonial even if it only sometimes breeds colonially, and as a cavity nester even if it sometimes lays in the open. RESULTS AND DISCUSSION Intraspecific nest parasitism was reported in 234 species of birds: one Struthioniformes, two Tinamiformes, two Procellariiformes, six Podicipediformes, five Ciconiiformes, one Phoenicopteriformes, 74 Anseriformes, one Falconiformes, 32 Galliformes, eight Gruiformes, 19 Charadriiformes, nine Columbiformes, five Cuculiformes, two Apodiformes, one Coraciiformes and 66 Passeriformes. INP and mode of reproduction As expected, INP rate was highest among precocial

134

Y Yom-Tov

birds, and was reported for most precocial orders. Among the 161 species of Anseriformes (number of species per order from Gill 19943, INP was reported in 74 species (46%). These INP rates are probably underestimates, as can be deduced from the following facts. Reported rates of INP are higher in areas where most ornithologists work: INP was reported in 46 species (70%) of the 66 species breeding in the western Palearctic and North America (USA and Canada), and in 16 species (70%) of the 23 species breeding in Australia and New Zealand. Hence, it is logical to expect that INP will be reported for yet more species of Anseriformes, as predicted by Sayler (1992). High rates of INP (but much lower than for Anseriformes) were reported for other precocial orders, including the Podicipediformes (29"L)), Galliformes (1 2%), Charadriiformes (5%) and Gruiformes (4%). O n the other hand, low INP rates were reported for altricial birds: 5% Ciconiiformes, 1Yo among Passeriformes and

even less among other orders. For some altricial orders it was not reported at all. For example, there are no reports of INP among the 358 Psittaciformes and 178 Strigiformes, only two species with INP in the Apodiformes (422 species), and one each of Falconiformes (3 11 species) and Coraciiformes (21 8 species).

INP and coloniality Only about 13% of bird species are colonial breeders (Hoyo et al. 1992) but out of the 232 species in Table 1 , 134 (57.5%) are colonial breeders. Among altricials exhibiting INP, 70% of those that are Passeriformes, all that are Ciconiiformes, the two species that are Procellariiformes, and t h e single species of Coraciiformes and Falconiformes are colonial. Among the precocial and semi-precocial Anseriformes (74 species; Table l ) , Charadriiformes (19 species) and Podicipediformes (six species), 44 (59"/0), 15 (79'Yo)

Table 1. The occurrence of intraspecific nest parasitism in 234 species of birds. The list is organized by order and family (following Voous 1985) and within each family by the alphabetical order of the genera. Coloniality is expressed as y (yes) or n (no), mode of reproduction as altricial (A), precocial (P) or semi-precocial (SP), nest type as open, closed and cavity. For most species only one reference is given (preferably the earliest report), although for some there are several. All studies apart from Victoria (1972) are field studies.

Species

Family

Altriciali Precocial

Colonial ~~

Nest type

Reference

~

Struthio camelus Ostrich

Struthionidae

n

P

Open

Bertram 1979

Nothura maculosa Spotted Tinamou Nothura darwinii Darwin's Tinarnou

Tinamidae Tinamidae

n n

P P

Open Open

Bump & Bump 1969 Bump & Bump 1969

Diomedea nigripes Black-footed Albatross Diomedea immutabilis Laysan Albatross

Diomedeidae Diomedeidae

Y

v

A A

Open Open

Tickell & Pinder 1966 Tickell & Pinder 1966

Aechmophorus clarkii Clark's Grebe Aechmophorus occidentalis Western Grebe fodiceps cristatus Great-crested Grebe Podiceps grisegena Red-necked Grebe fodiceps nigricollis Eared Grebe Podiceps occipitalis Silvery Grebe

Podicipedidae Podicipedidae Podicipedidae Podicipedidae Podicipedidae Podicipedidae

y y yin y/n y y

P P P P P P

Open Open Open Open Open Open

Storer & Nuechterlein 1992 Storer & Nuechterlein 1992 Cramp & Simmons 1977 Cramp & Simmons 1977 Lyon & Everding 1996 Nuechterlein in Rohwer & Freeman 1989

Ardea purpurea Purple Heron Ardeola ralloides Squacco Heron Egretta rufescens Reddish Egret

Ardeidae Ardeidae Ardeidae

Y Y Y

A A A

Open Open Open

Ciconia maguan Maguari Stork Eudocimus albus White Ibis

Ciconiidae Threskiornithidae

Y y

A

A

Open Open

Gonzalez-Martin & Ruiz 1996 Gonzalez-Martin & Ruiz 1996 Paul in Rohwer & Freeman 1989 Thomas 1984 Frederick 8, Shields 1986

Phoenicopterus ruber Greater Flamingo

Phoenicopteridae

y

P

Open

Brown 1958

Aix galericulata Mandarin Duck Aix sponsa Wood Duck

Anatidae Anatidae

n n

P

P

Cavity Cavity

Alopochen aegyptiacus Egyptian Goose

Anatidae

n

P

Cavity

Davies & Baggott 1989 Leopold 1951, Semel & Sherman 1992 Pitrnan 1965 continued

02001 British Ornithologists'Union, lba.

143, 133-143

lntraspecific nest parasitism

135

Table 1 continued.

Colonial

Altriciall Precocial

Species

Family

Anas castanea Chestnut Teal Anas clypeata Northern Shoveler Anas fulvigula Mottled Duck Anas gracilis Grey Teal Anas penelope Wigeon Anas plafyrhynchos Mallard

Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae

Anas querquedula Garganey Anas rubripes American Black Duck Anas strepera Gadwall Anas superciliosa Grey Duck Anser anser Greylag Goose Anser canagica Emperor Goose Anser caerulescens Snow Goose Anser erythropus Lesser White-fronted Goose Anser indicus Bar-headed Goose Anser rossi Ross Goose Anseranas semipalamata Magpie Goose Aythya affinis Lesser Scaup Aythya americana Redhead Aythya australis Australian White-eye Aythya ferina Pochard Aythya fuligula Tufted Duck Aythya marila Scaup Aythya nyroca Ferruginous White-eye Aythya valisineria Canvasback Biziura lobata Musk Duck Branta bernicla Brant Goose Branta canadensis Canada Goose Branta leucopsis Barnacle Goose

Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae

P P P P P P P P P P P P P P P P P P

Bucephala albeola Bufflehead Bucephala clangula Goldeneye

Anatidae Anatidae

P P

Bucephala islandica Barrow's Goldeneye Cairina moschata Muscovy Duck Chenonettajubata Maned Goose Clangula hyemalis Long-tailed Duck

Anatidae Anatidae Anatidae Anatidae

P P P P

Cygnus atratus Black Swan Dendrocygna autumnalis Black-bellied Whistling-duck Dendrocygna bicolor Fulvous Whist Iing-duck

Anatidae Anatidae

P P

Frith 1967 Dementiev et a/. 1967 Johnson etal. 1996 Frith 1967 Bengston 1972b Cramp & Simmons 1977, Evans & Williams 1987 Dementiev et a/. 1967 Open Open Stotts & Davis 1960 Duebbert 1966 Open Attiwill etal. 1981 Cavity Cramp & Simmons 1977 Open Open Eisenhauer & Kirkpatrick 1977 Barry 1960, Quinn et a/. 1987 Open Tegelstrom & Vonessen 1996 Open Open Weigmann & Lamprecht 1990 Bellrose 1976 Open Open Whitehead & Tschirner 1991 Open Bent 1902 Open Joyner 1983, Weller 1959 Frith 1967 Open Open Cramp & Simmons 1977 Newton & Campbell 1975 Open Bengston 1972b Open Dementiev et a/. 1967 Open Sorenson 1993 Open Attiwill etal. 1981 Open Barry 1960 Open Kossack 1950 Open Open Choudhury etal. 1993, Forslund & Larsson 1995 Phillips 1923-1926 Cavity Brestwer 1900, Dow & Cavity Fredga 1984 Openlcavity Bengston 1972b Philips 1922-1926 Cavity Briggs 1991 Cavity Dementiev etal. 1967, Open Alison 1975 Frith 1967 Open Cavity Bolen 1967

Anatidae

P

Cavity

Dendrocygna eytoni Plumed Whistling-duck Dendrocygna javanica Lesser Whistling-duck Dendrocygna viduata White-faced Whistling-duck Histrionicus histrionicus Harlequin Duck Lophodytes cucullatus Hooded Merganser Malacorhynchus membranaceus Pink-eared Duck Marmaronetta angustirostris Marbled Teal Melanitta fusca White-winged Scoter Melanitta nigra Black Scoter Mergus albellus Smew Mergus merganser Goosander

Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae

P P P P P P P P P P P

-

P P P P P P P P P P P

Nest type

Reference

Openlcavity Open Open Openlcavity Open Open

Meanley & Meanley 1959, Barnhart 1901 Frith 1967 Cavity Cavity Johnsgard 1978 Clark 1976 Cavity Bengston 1972a Open Morse & Wight 1969 Cavity Open Frith 1967 Open Hawkes 1970 Verneer 1968 Open Open Bengston 1972a Cramp & Simmons 1977 Open Openlcavity Cramp & Simmons 1977 continued

@ 2001 British Ornithologists' Union, Ibis, 143, 133-143

136

Y. Yom-Tov

Table 1 continued.

Species

Family ~

Colonial

~-

Altriciali Precocial

Nest type

Reference ~______

~~

Mergus serrator Red-breasted Merganser Neffa erythropthalma Southern Pochard Netta peposaca Rosybill Neffa rufina Red-crested Pochard Nettapus coromandehanus Cotton Pygmy Goose Oxyura australis Australian Blue-billed Duck Oxyura dominica Masked Duck Oxyura jamarcensis Ruddy Duck Oxyura leucocephala White-headed Duck Oxyura maccoa Maccoa Duck Oxyura vittata Argentine Blue-billed Duck Plectopterus gambensis Spur-winged Goose Sark/diornis melanotos Comb Duck Somateria moll/ssima Eider

Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae

Yln n Y/n Y/n n n n n n Y/n Y/n n Yln Y

P P P P P P P P P P P

P P P

Open Open Open Open Cavity Open Open Open Open Open Open Openkavity Openkavit y Open

Somateria spectabilis King Eider Sfictoneffa naevosa Freckled Duck Tadorna ferruginea Ruddy Shelduck Tadorna radjah Radjah Shelduck Tadorna fadorna Shelduck Tadorna tadornoides Australian Shelduck Tadorna variegata New Zealand Duck Thalassornis leuconofus White-backed Duck

Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae Anatidae

n n Y/n n Y n n Y/n

P P P P P P P P

Open Open Open Cavity Openlcavity Openkavity Cavity Open

Young & Titman 1987 Clancey 1967 Gibson 1920 Hellebrekers & Voous 1964 Ah & Ripley 1968-1974 Serventy & Whittell 1967 Bond 1961 Low 1941, Joyner 1983 Dementiev et a / 1967 Clark 1964 Gibson 1920 Pitman 1965 Pitman 1965 Erikstad e f a/ 1998, Robertson 1998 Cramp & Simmons 1977 Rogers 1990 Cramp & Simmons 1977 O'Brien 1990 Hori 1964, 1969 Frith 1967 Williams 1979 Clancey 1967

Falco naumanni Lesser Kestrel

Falconidae

Y

A

Open

Negro et a/. 1996

Orfalis vetula Plain Chachalaca Cracidae n Numida meleagris Crowned Guineafowl Numididae n Perdix perdix Gray Partridge Numididae n Francolinus africanus Greywing Francolin Numididae n Francolinus capensis Cape Francolin Numididae n Callipepla californica California Quail Odontophoridae n Callipepla douglasii Elegant Quail Odontophoridae n Colinus virginianus Bobwhite Quail Odontophoridae n Oreortyx pictus Mountain Quail Odontophoridae n Alectoris barbara Barbary Partridge Phasianidae n Alectoris chukar Chukar Partridge Phasianidae n Alectoris graeca Rock Partridge Phasianidae n Ammoperdix griseogularis See-see Partridge Phasianidae n Amrnoperdix heyi Sand Partridge Phasianidae n Coturnix corornandelica Black-breasted Quail Phasianidae n Coturnix coturnix Eurasian Quail Phasianidae n Coturnix delegorguei Harlequin QuaiI Phasianidae n Dendrortyx macroura Long-tailed Wood Partridge Phasianidae n Francolinus leucoscepus Yellow-necked Francolin Phasianidae n Francolinus natalensis Natal Francolin Phasianidae n Francolinus schlegelii Schlegel's Francolin Phasianidae n Guffera edouardi Crested Guineafowl Phasianidae n Lophura edwardsi Edward's Pheasant Phasianidae n Meleagris gallopavo Wild Turkey Phasianidae n Phasianus colchicus Ring-necked Pheasant Phasianidae n Tetraogallus caucasicus Caucasian Snowcock Phasianidae n Tympanuchus phasianellus Sharp-tailed Grouse Phasianidae n Lagopus lagopus Willow Ptarmigan Tetraonidae n

P P P P P P P P P P P P P P P P P P P P P P P P P P P P

Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open Open

Fleetwood & Bolen 1965 Clancey 1967 Knott etal. 1943 Clancey 1967 Clancey 1967 Tyler 1913 Johnsgard 1988 Klimstra & Roseberry 1975 Grinnell etal. 1918 Urban eta/. 1986 Mackie & Buechner 1963 Johnsgard 1988 Cramp & Simmons 1980 Yom-Tov 1980a Johnsgard 1988 Cramp & Simmons 1980 Urban e t a / . 1986 Johnsgard 1988 Urban et a/. 1986 Urban et a/. 1986 Johnsgard 1988 Clancey 1967 Schorger 1966 Johnsgard 1986 Knott etal. 1943 Dementiev et a/. 1967 Graison 1989 Martin 1984, Freeland e f a/. 1995 continued

@ 2001 British Ornithologists' Union, Ibis, 143, 133-143

lntraspecific nest parasitism

137

Table 1 continued.

Species

Family

Colonial

-

Altriciall Precocial

Nest type

Reference

~

Lagopus leucurus White-tailed Ptarmigan Dendrogapus canadensis Spruce Grouse Dendrogapus obscurus Blue Grouse Tetra0 tetrix Capercaillie

Tetraonidae Tetraonidae Tetraonidae Tetraonidae

n n n n

P P P P

Open Open Open Open

Choate 1963 Rand 1947 Redfield 1973 Cramp & Simmons 1980

Grus canadensis Sandhill Crane Amaurornis flavirostris Black Crake Crex crex Corn Crake Fulica americana American Coot Fulica atra Eurasian Coot Fulica cristata Crested Coot Gallinula chloropus Moorhen

Gruidae Rallidae Rallidae Rallidae Rallidae Rallidae Rallidae

n n n yln Y/n n Y/n

P P P P P P P

Open Open Open Open Open Open Open

Porphyrio porphyrio Green-backed Gallinule

Rallidae

Y

P

Open

Littlefield 1981 Urban et a/. 1986 Brown 1938 Arnold 1987, Lyon 1993 Horsfall 1984 Urban et a/. 1986 Gibbons 1986, McRae 1996 Craig 1980

Charadrius melodus Piping Plover Charadrius montanus Mountain Plover Limosa fedoa Marbled Godwit Limosa limosa Black-tailed Godwit Phalaropus fulicaria Red Phalarope Phalaropus tricolor Wilson’s Phalarope Vanellus vanellus Northern Lapwing Larus argentatus Herring Gull

Charadriidae Charadriidae Charadriidae Charadriidae Charadriidae Charadriidae Charadriidae Laridae

Y/n n Y Y/n yln n Yln Y

SP SP SP SP SP SP SP SP

Open Open Open Open Open Open Open Open

Larus delawarensis Ring-billed Gull Larus minutus Little Gull Sterna caspia Caspian Tern

Laridae Laridae Laridae

Y Y Y

SP SP SP

Open Open Open

Sterna dougallii Roseate Tern Sterna hirundo Common Tern

Laridae Laridae

Y Y

SP SP

Open Open

Rynchops niger Black Skimmer Catoptrophoryssemipalmatus Western Willet Recurvirostra americana American Avocet Scolopax minor American Woodcock Scolopax rusticola Eurasian Woodcock Catharacta lonnbergi Brown Skua

Rhynchopidae Scolopacidae Scolopacidae Scolopacidae Scolopacidae Stercoraiidae

Y Y Y n n

SP SP SP SP SP SP

Open Open Open Open Open Open

Hussell & Woodford 1965 Hamas & Graul 1985 Colwell 1986 Haverschmidt 1963 Kistchinski 1975 Colwell 1986 Spencer 1953 Burger in Rohwer & Freeman 1989 Conover et a/. 1979 Haverschmidt 1946 Cuthbert in Rohwer & Freeman 1989 Shealer & Zurovchak 1995 Nisbet in Rohwer & Freeman 1989 Erwin 1977 Colwell 1986 Giroux 1985 Lincoln 1951 Alexander 1946 Millar eta/. 1994

Columba livia Rock Dove Columba palurnbus Wood Pigeon Columba oenas Stock Dove Streptopelia risoria Ringed Turtle Dove Streptopelia senegalensis Palm Dove Streptopelia turtur Turtle Dove Streptopelia capiola Ring-necked Dove Oena capensis Namaqua Dove Zenaida macroura Mourning Dove

Columbidae Columbidae Columbidae Columbidae Columbidae Columbidae Columbidae Columbidae Columbidae

Y n n n Y/n y/n n n n

A A A A A A A A A

Open Open Open Open Open Open Open Open Open

Goodwin 1977 Murton 1965 Campbell in Cramp 1985 Goodwin 1977 Shoham 1975 Shoham 1975 Urban et a/. 1986 Urban et a/. 1986 Weeks 1980

Coccyzus americanus Yellow-billed Cuckoo

Coccyzidae

n

A

Open

Coccyzus eryfhrophthalmus Black-biIled Cuckoo Coccyzus melacoryphus Dark-billed Cuckoo Coccyzus pumilus Dwarf Cuckoo Geococcyx californianus Roadrunner Glaucis hirsufa Rufous-breasted Hermit

Coccyzidae Coccyzidae Coccyzidae Crotophagidae Trochilidae

n n n n n

A A A A A

Open Open Open Open Open

Nolan & Thompson 1975, Fleischer et a/. 1985 Baicich & Harrison 1997 Sick 1993 Ralph 1975 Miller 1946 Haverschmidt 1968

Y

~~

~

continued

@ 2001 British Ornithologists’ Union, lbrs, 143, 133-143

138

Y Yom-Tov

Table 1 continued.

Species

Family ~~~~~~

~

Colonial ~~

Altriciall Precocial

Nest type

~

Reference ~~

Threnetes rucken Band-tailed Barbthroat

Trochilidae

n

A

Open

Skutch 1973

Merops bullockoides White-fronted Bee-eater

Meropidae

Y

A

Cavity

Ernlen & Wrege 1986, Wrege & Ernlen 1987

Calocifta formosa White-throated Magpie-jay Gymnorhynus cyanocephalus Piiion Jay Pica pica Black-billed Magpie Passerculus sandwichensis Savannah Sparrow

Corvidae Corvidae Corvidae Emberizidae

n Y Y yJn

A A A A

Open Open Closed Open

Spizella passerina Chipping Sparrow

Ernberizidae

n

A

open

Spizella pusilla Field Sparrow Amandava amandava Red Munia Es trilda paludicola Fawn- b reasted Wax biII Lonchura bicolor Blue-billed Mannikin Lonchura cantans African Silverbill Lonchura kelaarfi Hill Munia Lonchura malabarica White-throated Munia Lonchura puncfulafa Spice Finch Taeniopygia guffata Zebra Finch Carduelis chloris European Greenfinch Carduelis tristis American Goldfinch

Ernberizidae Estrildidae Estrildidae Estrildidae Estrildidae Estrildidae Estrildidae Estrildidae Estrildidae Fringillidae Fringillidae

n Y Y Y Y Y Y Y Y yln n

A A A A A A A A A A A

open Closed Closed Closed Closed Closed Closedkavity closed open Open Open

Hirundo pyrrhonota Cliff Swallow

Hirundinidae

Y

A

Closed

Hirundo rustica Barn Swallow Hirundo spilodera South African Cliff Swallow Pefrochelidon ariel Fairy Martin Progne subis Purple Martin Riparia riparia Bank Swallow Tachycineta bicolor Tree Swallow

Hirundinidae Hirundinidae Hirundinidae Hirundinidae Hirundinidae Hirundinidae

Y Y Y Y Y

Y

A A A A A A

Open Closed Closed closed cavity Cavity

Agelaius phoeniceus Red-winged Blackbird

lcteridae

yJn

A

Open

Agelaius tricolor Tricolored Blackbird Dolichonyx oryzivorus Bobolink Euphagus cyanocephalus Brewer's Blackbird

lcteridae lcteridae lcteridae

Y Y

A A A

Open Open Open

Quiscalus quiscula Common Grackle

lcteridae

Y

A

closed

Xanthocephalus xanthocephalus Yellow-headed Blackbird Mimus polyglottos Northern Mockingbird

lcteridae

n

A

Open

Mirnidae

n

A

Closed

Anthus spinoleffa Alpine Water Pipit Ficedula hypoleuca Pied Flycatcher

Motacillidae Muscicapidae

n n

A A

Open Cavity

Nectarinia osea Orange-tufted Sunbird Parus major Great Tit Parus monfanus Willow Tit Dendroica petechia Yellow Warbler Passer domesficus House Sparrow

Nectarinidae Paridae Paridae Parulidae Passeridae

n n n n

A

Y

A A A A

Closed Cavity Cavity Open Closedlcavity

Passer melanurus Cape Sparrow

Passeridae

Y

A

Closed

Langen1996 Trost & Webb 1986 Trost & Webb 1986 Weatherhead & Robertson 1978 Middleton in Rohwer & Freeman 1989 Petter et a/. 1990 Goodwin 1982 Goodwin 1982 Goodwin 1982 Goodwin 1982 Goodwin 1982 Dhindsa 1983b Goodwin 1982 Birkhead et a/. 1990 Monk 1954 Middleton in Rohwer & Freeman 1989 Brown & Brown 1988, Smyth etal. 1993 M ~ l l e r1987 Earle 1986 Manwell & Baker 1975 Morton etal. 1990 Hoogland & Sherman 1976 Lombard0 1988, Lifjeld eta/. 1993 Orians in Rohwer & Freeman 1989 Emlen 1941 Gavin & Bollinger 1985 Orians in Rohwer & Freeman 1989 Howe in Rohwer & Freeman 1989 Harms et a/. 1991, Lyon et a/. 1992 Derrickson in Rohwer & Freeman 1989 Reyer ef a/. 1997 G. Hogstedt in Haland 1986, Gelter & Tegelstrom 1992 Zilberman eta/. 1999 Verboven & Mateman 1997 Orell etal. 1997 Sealy etal. 1989 Burke & Bruford 1987, Wetton e t a / . 1987 Earle 1986

n

continued

@ ZOO1 British Ornithologists' Union, Ibis, 143, 133-143

lntraspecific nest parasitism

139

Table 1 continued

Species

Family

Colonial

Y

Altriciall Precocial

Nest type Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Closed Cavity Cavity Cavity Cavity

cavity cavity Open Open Open

Passer moabificus Dead Sea Sparrow Passer monfanus Tree Sparrow Euplecfes afer Golden Bishop Euplecfes axillaris Red-shouldered Widow Euplecfes capensis Yellow-rumped Bishop Euplecfes orix Red Bishop Ploceus baglafecht Baglafecht Weaver Ploceus benghalensis Indian Weaverbird Ploceus bojeri Golden Palm Weaver Ploceus cucullafus Village Weaver Ploceus intermedius Lesser-masked Weaver Ploceus manyar Striated Weaver Ploceus nigerrimus Vieillot's Black Weaver Ploceus ocularis Spectacled Weaver Ploceus philippinus Baya Weaverbird Ploceus rubiginosus Chestnut Weaver Ploceus faeniopferus Northern-masked Weaver Ploceus velafus Southern-masked Weaver Ploceus xanfhops Holub's Golden Weaver Sturnus cineraceus Grey Starling Sfurnus roseus Rose-coloured Starling Sfurnus unicolor Spotless Starling Sfurnus vulgaris European Starling

Passeridae Passeridae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Ploceidae Sturnidae Sturnidae Sturnidae Sturnidae

Y Y Y Y/n Y Y y/n

A A A A A A A A A A A A A A A A A A A A A A A

Jroglodytes aedon House Wren Siala sialis Eastern Bluebird Jurdus grayi Clay-colored Robin Turdus migraforius American Robin Jurdus pilaris Fieldfare

Troglodytidae Turdidae Turdidae Turdidae Turdidae

n n n n n

A A A A A

and 6 (100%) species, respectively, are colonial. Evidently, INP is closely related to coloniality. In contrast, none of the Galliformes is colonial, hence there is a relatively low rate of INP (12%) in this precocial order. In summary, high rates of INP exist among precocial birds and are associated with coloniality, as reported earlier (Yom-Tov 1980a, Andersson 1982, Rohwer & Freeman 1989).

NOTE ADDED IN PROOF After the acceptance of this paper, intraspecific nest parasitism was reported in two more avian species: the Pied Avocet Recuwirostru uvosettu (Hotker 2000) and the Emu Drornaius novaehollandiue (Taylor et al. 2000). These additions raise the number of species for which INP is reported to 236. I am grateful to Jon Wright for encouragement and hospitality, to Robert Payne, Harry Power and an anonymous referee

Y Y Y Y Y Y Y Y Y

Y Y Y Y Y

Y

Reference Yom-Tov 1980b Summers-Smith 1988 Maclean 1993 Maclean 1993 Maclean 1993 Maclean 1993 Freeman 1988 Dhindsa 1983a Freeman 1988 Victoria 1972 Freeman 1988 Dhindsa 1983a Freeman 1988 Freeman 1988 Dhindsa 1990 Freeman 1988 Jackson 1992 Freeman 1988 Freeman 1988 Yamaguchi & Saitou 1997 Cramp & Perrins 1994 Cramp & Perrins 1994 Yom-Tov ef a/. 1974, Evans 1988, Romagnano et a/. 1990 Picman & Belles-Isles 1988 Gowaty & Karlin 1984 Dryrez 1983 Gowaty & Davies 1986 Haland 1986

for their comments, to Arieh Landsman for his help with compiling the data from the literature and to Mary Fox and Dorothy Vincent for careful editing. This work was supported by the Israel Cohen Chair for Environmental Zoology.

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