We summarized 40 reports of nine species of brood parasites feeding young of their own species. These observations suggest that the propensity to provision young has not been lost entirely in brood parasites despite the belief that brood parasitic adults abandon their offspring at the time of laying. The hypothesis that species that participate in courtship feeding are more likely to provision young was not supported: provisioning of young has been observed in two species of brood parasites that do not courtship feed. The function of this provisioning is unknown, but we suggest it may be: (1) a non-adaptive vestigial behavior or (2) an adaptation to ensure adequate care of parasitic young. The former is more likely the case. Further studies are required to determine whether parasitic adults commonly feed their genetic offspring.

Department of Zoology University of Manitoba Winnipeg, Manitoba R3T 2N2 Canada ADULTOS DE AVES PARASTICAS ALIMENTANDO PICHONES Y VOLANTONES DE SU PROPIA ESPECIE: UNA REVISION Sinopsis.--Resumimos 40 informes de nueve especies de aves parasiticas que alimenaron a pichones de su propia especie. Las observaciones sugieren que la propensividad de alimentar a los pichones no ha sido totalmente perdida en las aves parasiticas, no empece a la creencia de que los parasiticos abandonan su progenie al momento de poner los huevos. La hipttesis de que las especies que participan en cortejo de alimentacitn, son mils propensas a alimentar los pichones no tuvo apoyo. Las observaciones de alimentacitn a pichones se han hecho en dos especies parasiticas cuyo cortejo no incluye la alimentacitn de la pareja. La funcitn de proveer alimento se desconoce. No obstante, sugerimos que pueda ser: 1) una conducta vestigial no adaptativa, o 2) una adaptacitn parc asegurar el cuidado adecuado de los pi- chones parasiticos. E1 61timo parece ser la causa mgts probable. Se necesitan estudios mils detaliados parc determinar si los adultos de especies parasiticas alimentan a sus pichones comfnmente. Arian brood parasitism is a reproductive strategy that has evolved in 1% of all bird species. Obligate brood parasites lay all of their eggs in nests of other species and are believed to provide no parental care to their offspring. They leave the incubation of their eggs and feeding of their offspring to hosts and do not normally participate in these behaviors (Payne 1977). Interestingly, it has come to our attention that some brood parasites have been observed feeding young of their own species. We reviewed the literature and compiled observations of this behavior (1) to determine which parasitic species have been observed provisioning young; (2) to determine how frequently this behavior has been described in the literature; (3) to describe the nature of the observations (i.e., the sex of the provisioner, whether nestlings or fledglings are involved, and duration of the feeding); and (4) to identify the possible function of this behavior. TREATMENT OF RECORDS Published observations were mainly anecdotal because it was not known in any case whether the provisioning adults were the genetic parents of the young that were fed. Care must be taken when interpreting the ap- parent feeding of a young bird because some observers may have mistook it for feeding of an adult female, which occurs during courtship feeding. We determined, therefore, which species also are known to practice court- ship feeding because we wanted to determine whether the records of provisioning involved only these species. Species that courtship feed may be more predisposed to feeding young (Benson and Serventy 1957). If the provisioning of young is an extension of courtship feeding, then only males would be expected to perform this behavior because only they feed females. Common names of species were used throughout the paper (sci- entific names in Appendix); taxonomy follows Sibley and Monroe (1990). OBSERVATIONS OF PROVISIONING OF YOUNG BY BROOD PARASITES Feeding of young brood parasites by individuals of the same species has been reported at least 40 times (Table 1). This behavior has been observed in at least nine parasitic species from the families Cuculidae and Fringillidae (Tribe Icterini), and has been reported most often in Pallid (n = 9) and Dideric (n = 11) Cuckoos (Table 1). Other authors have made only a general statement that the following parasites feed young of their own species, but did not provide details of the observations: Pied Cuckoo (Ali and Ripley 1969), Fan-tailed Cuckoo (Friedmann 1968), Shining Bronze-cuckoo (Oliver 1955), Dideric Cuckoo (Mackworth-Praed and Grant 1970, Roberts 1958), Channel-billed Cuckoo (Friedmann 1968), and cuckoos in general (Chisholm 1956, Fletcher 1915). Including these observations, provisioning of young by a brood parasite has been recorded in at least 12 parasitic species. Ten of these species also court- ship feed. Thus, "parental" feeding has been observed in two species that do not courtship feed: Asian Koel and Brown-headed Cowbird (Table 2). Species that courtship feed are not more likely to provision young (Fisher exact test, one-tailed, P > 0.05). Five reports involved nestlings and 38 involved fledglings (three reports involved both nestlings and fledglings). Most authors did not report the sex of the individual that fed the young, but in those cases where birds were sexed, provisioners included males (n = 7), females (n = 6), a pair of adults (n = 3), and one group of adults of both sexes. A pair was not assumed to be male and female. Some adult parasites were observed feed- ing more than one parasitic young (Baird 1945, Merritt 1956). Some observations were made over several days (J. A. Fletcher 1915, L. R. Fletcher 1925, Hanscombe 1915, Hare 1915, Hume in Fulton 1904, Kik- kawa and Dwyer 1962, Van Someren 1956, Walton 1903); however, be- cause the individuals were not banded, their identities were not known. McCracken (1984) "believed" a male Bronzed Cowbird fed one of three cowbird fledglings that was in its company, but the evidence was insufficient to include in Table 1. The following records were also ex- cluded from our review because they were criticized: (1) Cheeseman (1890) and (2) Fulton (1910) were criticized by Friedmann (1949), (3) Graham (1940) by Watson and Bull (1950), (4) Hartlaub (in Friedmann TABLE 1. Feeding of parasitic nestlings or fledglings by an adult of the same species. Known or suspected Parasite species host species a Comments Reference Great Spotted Artamus sp. (s) adult fed fledgling North 1912 Cuckoo Great Spotted U Cuckoo Common Cuckoo Common Cuckoo Common Cuckoo Common Cuckoo U Pallid Cuckoo Pallid Cuckoo Pallid Cuckoo adult fed "juvenile" U 57 fed recent fledgling U adult fed fledgling Meadow-pipit (k) more than one observation of adult feeding cuckoo during nestling and fledg- ling stages adult fed young U adult fed young (2 reports) Artamus sp. (s) adult fed fledgling U Pallid Cuckoo U Pallid Cuckoo U Pallid Cuckoo U Pallid Cuckoo U Pallid Cuckoo U Pallid Cuckoo Pallid Cuckoo or Shining Bronze- cuckoo Shining Bronze- cuckoo Klaas's Cuckoo Klaas's Cuckoo Klaas's Cuckoo Klaas's Cuckoo Klaas's Cuckoo African Emerald Cuckoo Dideric Cuckoo White-browed Scrubwren (k) Yellow-rumped Thornbill (k) Grey Gerygone (s) u sparrow (Passer sp.) (k) Village Weaver (s) Dideric Guckoo Dideric Cuckoo Mundy and Cook 1977 Browne 1886 Kelin 1911 Bannerman and Lodge 1955:132 Palmer in Bannerman and Lodge 1955: 132 Campbell 1900 North 1912 pair of cuckoos assisted host Hanscombe 1915 daily in feeding fledgling adult fed "well-grown Jackson 1949 young" adult twice fed young adult cuckoo fed fledgling adult fed fledgling; another adult fed a different fledg- ling repeatedly during one period of observation feeding young or courtship Fleming 1979 feeding of immature 57 57 fed fledgling Ambrose 1987 Learmonth 1949 Cooper 1958 Kikkawa and Dwyer 1962 adult fed nestling Howe 1905 adult fed nearly full grown Hursthouse 1944 cuckoo adult fed young (4 reports), Moreau and Moreau one young not able to fly 1939 adult fed young (3 reports) Van Someren 1939 adult fed fledgling Moreau 1944 observations of cs feeding Van Someren 1956 nestlings and fledglings on separate occasions c fed 2 young repeatedly for Baird 1945 15 min c fed young Millar 1943 pair fed fledgling daily for a Hare 1915 week or more captive 5? fed captive fiedg- Millar 1926 ling; c fed same fledgling through cage bars 8 fed young or courtship Moreau 1944 feeding TABLE 1. Continued. [367 Known or suspected Parasite species host species Comments Reference Dideric Cuckoo U feeding young or courtship feeding Dideric Cuckoo U c fed "well-grown" young or courtship feeding Dideric Cuckoo U c fed young or courtship feeding observed 10 times in 6 min Dideric Cuckoo U on several separate occa- sions, different adults fed different young recently able to fly Dideric Cuckoo U c fed fully fledged young Dideric Cuckoo U Dideric Cuckoo U Dideric Cuckoo U Asian Koel U Brown-headed Cowbird Brown-headed Cowbird Brown-headed Cowbird Rose-breasted Grosbeak (k) adults fed fledglings on sev- eral occasions c fed young adult fed fledgling on several separate occa- sions, different adults fed different young 9 fed nestling daily Common Yellow- 9 fed cowbird at nestling throat (k) U Brown-headed U Cowbird Brown-headed Northern Cardi- Cowbird nal (s) and fledgling stage 9 fed juvenile repeatedly during one day of observa- tions within a flock, c c and 9 9 fed many young Bannerman in Mo- reau 1944 Friedmann 1948 Maclaren 1952 Maclaren 1953 Symons in Friedmann 1956 Smith 1957 Thomas 1960 Ottow and Duve 1965 Hume in Fulton 1904 Bonwell 1895 Walton 1903:219 Fletcher 1925 Merritt 1956 and 9 cowbird and 9 car- Hernandez 1986 dinal fed young cowbird Known host species (k), suspected host species (s) provided by the author, unknown host species (U). 1955) by Friedmann (1955), (5) Walton (1892) by Fletcher (1925), and (6) Worman (1930) by the editor of the Oologists' Record. Graham (1950) addressed Watson and Bull's (1950) criticisms. COURTSHIP FEEDING AND PROVISIONING OF YOUNG Observations of provisioning of young by brood parasites provide strong evidence that the propensity to provision young has not been lost by at least some individuals of some species of brood parasites. This be- havior is either vestigial or a new development (Kikkawa and Dwyer 1962). If the latter, this behavior may have developed as an extension of court- ship feeding (Benson and Serventy 1957). Courtship feeding and "pa- rental" feeding have similar elements, although the motives for these behaviors differ. Courtship feeding allows females to assess the food-gath- TAI.} 2. Parasitic species that have been observed provisioning young and/or courtship feeding. Observations of feeding Court- Parasitic species Young ship Courtship feeding references INDICATORIDAE Greater Honeyguide N a N CUCULIDAE Pied Cuckoo Y Y Great Spotted Cuckoo Y Y Thick-billed Cuckoo N Y Red-chested Cuckoo N Y Black Cuckoo N Y Common Cuckoo Y Y Pallid Cuckoo Y Y Brush Cuckoo N Y Fan-tailed Cuckoo Y Y Shining Bronze-cuckoo Y Y Horsfield's Bronze-cuckoo N Y Klaas's Cuckoo Y Y African Emerald Cuckoo Y Y Asian Koel Y N Long-tailed Koel N b N Channel-billed Cuckoo Y Y FRINGILLIDAE Bay-winged Cowbird N N Screaming Cowbird N N Shiny Cowbird N N Bronzed Cowbird N c N Brown-headed Cowbird Y N not mentioned in Friedmann 1955 d Godfrey 1939, Liversidge 1971 Channer 1976 Rowan 1983 Rowan 1983 Young 1946 Wyllie 1981 Klapste 1981, Lord 1956, Noske 1978, Robinson 1950, White 1950 Noske 1978 Chisholm 1940, Smithers 1977, Noske 1978 Falla et al. 1978, Serventy 1958, Wat- son and Bull 1950 McCulloch 1967, Moffat 1978, Wall 1978, Watson 1955 Winterbottom 1939 Haydock 1950 n/a n/a Goddard and Marchant 1983 not mentioned in Friedmann 1929 d not mentioned in Friedmann 1929 a not mentioned in Friedmann 1929 a not mentioned in Friedmann 1929 a not mentioned in Friedmann 1929 a a Possible observation by Hartlaub in Friedmann (1955:162). b Possible observation by Cheeseman (1890). c Possible observation by McCracken (1984). d Courtship feeding was not mentioned to occur in these species although there was a section on courtship behavior in Friedmann (1929, 1955). ering ability of males, increases the female's nutritional reserves, and fa- cilitates pair formation (Nisbet 1973, Smith 1980). "Parental" feeding, on the other hand, contributes primarily to the growth and development of the parasite. Courtship feeding, which also occurs in nonparasitic cuck- oos (e.g., Dwarf Cuckoo, Ralph 1975; Black Coucal, Rowan 1983; Black- billed Cuckoo, Spencer 1943) and many other nonparasitic taxa, has been reported in 11 avian orders and in almost three-quarters of passerine families (Smith 1980). Although provisioning of young occurs in many of the same parasitic species that also courtship feed, there is not enough evidence to suggest that the provisioning of young developed from court- ship feeding. First, both male and female brood parasites have been ob- served provisioning young, whereas only males feed adult females. Sec- ond, the provisioning of young has been observed in two parasitic species that do not courtship feed (Asian Koels and Brown-headed Cowbirds). Third, there is no significant association between courtship feeding and provisioning of young. It is not surprising that female brood parasites have been observed feeding nestlings or fledglings because they probably are the ones that find a nest and determine when it is suitable for laying and, in the case of many species, visit it to remove or pierce a host egg (e.g., Brooker and Brooker 1989, Carter 1986, Gill 1983, Jensen and Jensen 1969, Livesey 1936, Mason 1980, Sealy 1992). Males are not commonly involved in any of these activities, although there are a few observations of activity by parasitic males around nests (e.g., Sealy 1994). In light of the fact that parasitic males are seldom observed around host nests, it is interesting that males of the following species have been observed feeding nestlings and fledglings: Klaas's Cuckoo, African Emerald Cuckoo, Dideric Cuckoo, and Brown-headed Cowbird. PARENT-YOUNG ASSOCIATION Use of molecular genetic techniques would determine whether the adults are the genetic parents of the parasitic young they are feeding. Hahn and Fleischer (1995) found that female and juvenile Brown-headed Cowbirds trapped together had higher band similarities based on restric- tion fragments than would be expected by chance. This finding, however, was based on only 11 pairs of adult female and juvenile cowbirds, and individuals trapped at the same time may not have arrived at the trap together. Also, baited traps possibly affect the distribution of birds and may unnaturally attract many birds to a small area. Nonetheless, Hahn and Fleischer's finding is interesting because young cowbirds appeared to associate with their mothers. Fletcher (1925) observed an adult female cowbird that repeatedly fed the same juvenile although there were other young cowbirds feeding in the same area. The relationship between these individuals, however, was not known. Mother-offspring associations could result from female cowbirds re- cruiting their young, or from young cowbirds seeking out their mothers (McCracken 1984, Hahn and Fleischer 1995). The only way that young cowbirds could become visually familiar with their mothers is if females visited the nest during the nestling stage. Female cowbirds are known to visit nests after parasitism to remove host eggs (e.g., Mayfield 1961, Sealy 1992), however, mother-offspring associations cannot develop when nests are visited during the egg stage. Young cowbirds could become acousti- cally familiar with their mothers if the latter vocalize near the nest (Hahn and Fleischer 1995). Females frequently are in the area of a nest they have parasitized because there is evidence that they have home ranges (e.g., Alderson 1996, Darley 1983, Rothstein et al. 1984, Teather and Rob- ertson 1985). Hahn and Fleischer (1995) suggested that by associating with adult brood parasites, juveniles can better learn species-specific behavior. How- ever, species-specific behavior of brood parasites, such as singing in males and song recognition in females, has been found to be innate (King and West 1977), and Friedmann (1929) reported that large flocks of Brown- headed Cowbirds formed during the fall are frequently composed entirely of juveniles. Therefore, juvenile parasites likely seek out individuals of their own species rather than being recruited by adults. There is evidence that Great Spotted Cuckoos monitor nests after lay- ing. Soler et al. (1995) suggested that females of this species monitor nests they have parasitized to ensure that their egg remains in the nest. Nest predation, presumably by female cuckoos, is more likely to occur if the parasitic egg has been ejected. This forces hosts to renest and provides the cuckoos with another chance to lay an egg (see also Zahavi 1979). Similarly, Arcese et al. (1996) suggested that cowbirds depredate nests that are discovered too late in the host's nesting cycle for parasitism to succeed. Nest protection, another type of parental behavior, has been observed in Brown-headed Cowbirds and Great Spotted Cuckoos. Balda and Ca- rothers (1968) twice observed a female Brown-headed Cowbird alarm call and display at the approach of a potential predator to a parasitized nest, and Gabrielson (1921) described another instance of this behavior. Fe- males should benefit by monitoring the progress of a nest and protecting the nest from the approach of predators, but the costs of spending so much time in this activity, however, likely outweigh the benefits. The scar- city of reports of nest protection by female cowbirds suggests that this behavior is rare. The female cowbirds probably were not at the nest spe- cifically to defend it, but rather defended the nest opportunistically. On the other hand, nest protection by Great Spotted Cuckoos appears to be more common. Soler et al. (unpubl. data in Soler et al. 1995) were scold- ed by cuckoos on 25% of 56 visits to 30 parasitized Black-billed Magpie nests, whereas they were scolded by magpies on only 5% of the visits. STRATEGY OR OPPORTUNISM. > About 90% of the records of provisioning in brood parasites involved cuckoos. This may reflect the fact that courtship feeding is found only among the cuckoos. Evidence for this, however, is weak because we did not find a significant association between courtship feeding and provi- sioning behavior. More likely, it is because there are more species of par- asitic cuckoos than cowbirds and honeyguides. The question remains whether the provisioning of young by brood par- asites is common enough to be considered a strategy or whether it is simply trivial behavior. Like other species of birds, brood parasites may possess an innate response to feed begging young (see Craig and Jamie- son 1990). If parasitic adults respond to the stimulus of begging young, then there should also be observations of adult parasites feeding begging young of other species. We did not find observations of this behavior, although there are many cases reported of nonparasitic species feeding other nonparasitic species (Shy 1982), and individuals other than the original foster parents of parasitic species feeding parasitic young (Sealy and Lorenzana 1997). The lack of such observations suggests that it is the adult brood parasite's intention to feed an individual of its own species. Implicitly, the adults recognize young of their own species. Provisioning of young by brood parasites may be important if hosts cannot provide the young parasite with enough or the right kind of food. In such cases, limited parental care by brood parasites could be an ad- aptation to poor-quality hosts. Benson and Serventy (1957) suggested that provisioning by brood parasites may be necessary in cases where insectiv- orous brood parasites lay eggs in nests of herbivorous species. There is, however, no evidence for this because parasitic nestlings raised by herbiv- orous species usually do not survive to fledging (Eastzer et al. 1980, Kozlo- vic et al. 1996, Middleton 1991; but see Seel and Davis 1981). Further- more, most observations involve brood parasites feeding fledglings, not nestlings. Brood parasitic species should be strongly selected to avoid her- bivorous hosts altogether (Kozlovic et al. 1996), rather than to compen- sate for herbivorous hosts by feeding the young at the nest. ACKNOWLEDGMENTS This review was initiated during a sabbatical leave granted to SGS by the University of Manitoba. N. L. Sealy provided invaluable help with the early stages of the literature search. Comments from J. V. Briskie and C. R. Chandler greatly improved the manuscript. Financial support was provided by a NSERC research grant to SGS and NSERC and (University of Manitoba) Faculty of Science Undergraduate Summer Research awards to JCL. LITERATURE CITED ALDERSON, G. W. 1996. Molecular genetic analysis of the mating system and host choice of an obligate brood parasitic bird, the Brown-headed Cowbird (Molothrus ater). M.Sc. thesis. McMaster Univ., Hamilton, Ontario. ALI, S. A., do S. D. RIPLE3/4. 1969. Handbook of the birds of India and Pakistan. Vol. 3. Oxford Univ. Press, Bombay, 325 pp. AMBROSE, S.J. 1987. Adult Fan-tailed Cuckoo Cuculus pyrrohophanus feeds fledgling. Emu 87:69. ARCESE, P., J. N.M. SMITH, AND M. I. HATCH. 1996. 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Common and scientific names of avian species mentioned in the paper. a FAMILY Common name Scientific name INDICATORIDAE Greater Honeyguide CUCULIDAE Pied Cuckoo Great Spotted Cuckoo Thick-billed Cuckoo Red-chested Cuckoo Common Cuckoo Pallid Cuckoo Brush Cuckoo Fan-tailed Cuckoo Shining Bronze-cuckoo Klaas's Cuckoo African Emerald Cuckoo Dideric Cuckoo Asian Koel Long-tailed Koel Channel-billed Cuckoo CENTROPODIDAE Black Cuckoo COCCYZIDAE Dwarf Cuckoo Black-billed Cuckoo CORXqDAE Black-billed Magpie MELIPHAGIDAE White-plumed Honeyeater PARDALOTIDAE White-browed Scrubwren Striated Calamanthus Yellow-rumped Thornbill Grey Gerygone VIREONIDAE Red-eyed Vireo PASSERIDAE House Sparrow Meadow pipit Village Weaver FRINGILLIDAE Yellow Warbler Common Yellowthroat Rose-breasted Grosbeak Northern Cardinal Bay-winged Cowbird Screaming Cowbird Shiny Cowbird Bronzed Cowbird Brown-headed Cowbird Indicator indicator Oxylophus jacobinus Clamator glandarius Pachycoccyx audeberti Cuculus solitarius Ct. canor,rs Cu. pallidus Cacomantis variolosus Ca. flabelliformis Chrysococcyx lucidus Ch. klaas Ch. cupreus Ch. caprius Eudynamys scolopacea E. taitensis Scythrops novaehollandiae Centropus grillii Coccyzus pumilus Co. erythropthalmus Pica pica Lichenostomus penicillata Sericornis frontalis Calamanthus fuliginosus Acanthiza chrysorrhoa Gerygone igata Vireo olivaceus Passer domesticus Artthus pratensis Ploceus cucullatus Dendroica petechia Geothlypis trichas Pheucticus ludovicianus Cardinalis cardinalis Molothrus badius M. rufoaxillaris M. bonariensis M. aeneus M. ater Order and nomenclature follow Sibley and Monroe (1990).