J. Field Ornithol., 69(3):364-375

ADULT BROOD PARASITES FEEDING NESTLINGS AND FLEDGLINGS OF THEIR OWN : A REVIEW

JANICEC. LORENZANAAND SPENCER G. SEALY Departmentof Zoology Universityof Manitoba Winnipeg,Manitoba R3T 2N2 Canada Abstract.--We summarized 40 reports of nine speciesof brood parasitesfeeding young of their own species.These observationssuggest that the propensityto provisionyoung hasnot been lost entirely in brood parasitesdespite the belief that brood parasiticadults abandon their offspringat the time of laying.The hypothesisthat speciesthat participatein courtship feeding are more likely to provisionyoung was not supported:provisioning of young has been observedin two speciesof brood parasitesthat do not courtshipfeed. The function of this provisioningis unknown, but we suggestit may be: (1) a non-adaptivevestigial behavior or (2) an adaptation to ensure adequatecare of parasiticyoung. The former is more likely the case.Further studiesare required to determinewhether parasiticadults commonly feed their genetic offspring. ADULTOS DE AVES PARAS•TICASALIMENTANDO PICHONES Y VOLANTONES DE SU PROPIA ESPECIE: UNA REVISION Sinopsis.--Resumimos40 informes de nueve especiesde avesparasiticas que alimenaron a pichonesde su propia especie.Las observacionessugieren que la propensividadde alimentar a los pichonesno ha sido totalmente perdida en las avesparasiticas, no empecea la creencia de que los parasiticosabandonan su progenie al momento de poner los huevos.La hipttesis de que las especiesque participan en cortejo de alimentacitn, son milspropensas a alimentar los pichonesno tuvo apoyo.Las observacionesde alimentacitn a pichonesse han hecho en dos especiesparasiticas 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 vestigialno adaptativa,o 2) una adaptacitn parc asegurarel cuidado adecuadode los pi- chonesparasiticos. E1 61timo parece ser la causamgts probable. Se necesitanestudios mils detaliados parc determinar si los adultos de especiesparasiticas alimentan a sus pichones comfnmente.

Arian brood parasitism is a reproductive strategy that has evolved in 1% of all species.Obligate brood parasiteslay all of their eggsin nests of other speciesand are believed to provide no parental care to their offspring. They leave the incubation of their and feeding of their offspringto hostsand do not normally participatein thesebehaviors (Payne1977). Interestingly,it hascome to our attentionthat somebrood parasiteshave been observedfeeding young of their own species.We reviewedthe literature and compiled observationsof this behavior (1) to determine which parasitic species have been observed provisioning young; (2) to determine how frequentlythis behaviorhas been described in the literature; (3) to describe the nature of the observations (i.e., the sex of the provisioner,whether nestlingsor fledglings are involved,and duration of the feeding); and (4) to identify the possiblefunction of this behavior.

TREATMENT OF RECORDS Publishedobservations were mainly anecdotalbecause it wasnot known in any casewhether the provisioningadults were the genetic parents of

364 Vol.69, No..• BroodParasites Feeding Young [365 the young that were fed. Care must be taken when interpreting the ap- parent feeding of a young bird becausesome observers may havemistook it for feeding of an adult female, which occursduring courtshipfeeding. We determined, therefore, which speciesalso are known to practicecourt- ship feeding becausewe wanted to determine whether the records of provisioninginvolved only these species.Species that courtship feed may be more predisposedto feedingyoung (Bensonand Serventy1957). If the provisioningof young is an extensionof courtshipfeeding, then only maleswould be expected to perform this behaviorbecause only they feed females.Common namesof specieswere usedthroughout the paper (sci- entific namesin Appendix); taxonomyfollows Sibley and Monroe (1990).

OBSERVATIONS OF PROVISIONING OF YOUNG BY BROOD PARASITES Feeding of young brood parasitesby individuals of the same species has been reported at least 40 times (Table 1). This behavior has been observedin at leastnine parasiticspecies from the familiesCuculidae and Fringillidae (Tribe Icterini), and has been reported most often in Pallid (n = 9) and Dideric (n = 11) (Table 1). Other authors have made only a general statementthat the followingparasites feed young of their own species,but did not provide details of the observations:Pied (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 cuckoosin general (Chisholm 1956, Fletcher 1915). Including these observations,provisioning of young by a has been recorded in at least 12 parasiticspecies. Ten of these speciesalso court- ship feed. Thus, "parental" feeding has been observedin two speciesthat do not courtshipfeed: Asian and Brown-headedCowbird (Table 2). Speciesthat courtshipfeed are not more likely to provisionyoung (Fisher exact test, one-tailed, P > 0.05). Five reportsinvolved nestlings and 38 involvedfledglings (three reports involvedboth nestlingsand fledglings).Most authorsdid not report the sex of the individual that fed the young, but in those caseswhere 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 wasnot assumedto be male and female. Some adult parasiteswere observedfeed- ing more than one parasiticyoung (Baird 1945, Merritt 1956). Some observationswere 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 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 becausethey were criticized: (1) Cheeseman (1890) and (2) Fulton (1910) were criticized by Friedmann (1949), (3) Graham (1940) by Watsonand Bull (1950), (4) Hartlaub (in Friedmann 366] J. C. Lorenzanaand S.G. Sealy j. FieldOrnithol. Summer 1998

TABLE1. Feeding of parasiticnestlings or fledglingsby an adult of the same species.

Known or suspected Parasitespecies host speciesa Comments Reference Great Spotted Artamussp. (s) adult fed fledgling North 1912 Cuckoo Great Spotted U adult fed "juvenile" Mundy and Cook Cuckoo 1977 CommonCuckoo U 57fed recentfledgling Browne 1886 U adult fed fledgling Kelin 1911 Common Cuckoo Meadow-pipit(k) more than one observation Bannerman and of adult feeding cuckoo Lodge 1955:132 during nestlingand fledg- ling stages Common Cuckoo U adult fed young Palmer in Bannerman and Lodge 1955: 132 U adult fed young (2 reports) Campbell 1900 Pallid Cuckoo Artamus sp. (s) adult fed fledgling North 1912 Pallid Cuckoo U pair of cuckoosassisted host Hanscombe1915 daily in feeding fledgling Pallid Cuckoo U adult fed "well-grown Jackson1949 young" Pallid Cuckoo U adult twice fed young Learmonth 1949 Pallid Cuckoo U adult cuckoofed fledgling Cooper 1958 Pallid Cuckoo U adult fed fledgling; another Kikkawaand Dwyer adult fed a different fiedg- 1962 ling repeatedlyduring one period of observation Pallid Cuckoo U feedingyoung or courtship Fleming1979 feedingof immature 57 Pallid Cuckoo White-browed 57fed fledgling Ambrose1987 Scrubwren (k) Pallid Cuckoo or Yellow-rumped adult fed nestling Howe 1905 Shining Bronze- Thornbill (k) cuckoo Shining Bronze- Grey adult fed nearly full grown Hursthouse1944 cuckoo (s) cuckoo Klaas's Cuckoo u adult fed young (4 reports), Moreau and Moreau one youngnot able to fly 1939 Klaas's Cuckoo adult fed young (3 reports) Van Someren 1939 Klaas's Cuckoo adult fed fledgling Moreau 1944 Klaas's Cuckoo observationsof c•sfeeding Van Someren 1956 nestlingsand fledglingson separateoccasions Klaas's Cuckoo c• fed 2 youngrepeatedly for Baird 1945 15 min African Emerald c• fed young Millar 1943 Cuckoo Dideric Cuckoo sparrow pair fed fledglingdaily for a Hare 1915 (Passersp.) (k) week or more Dideric Guckoo Village Weaver captive5? fed captivefiedg- Millar 1926 (s) ling; c• fed samefledgling through cage bars Dideric Cuckoo 8 fed young or courtship Moreau 1944 feeding Vol. 69, No. 3 BroodParasites Feeding Young [367

TABLE 1. Continued.

Known or suspected Parasitespecies host species• Comments Reference Dideric Cuckoo U feeding young or courtship Bannerman in Mo- feeding reau 1944 Dideric Cuckoo U c• fed "well-grown"young Friedmann 1948 or courtshipfeeding Dideric Cuckoo U c• fed young or courtship Maclaren 1952 feeding observed10 times in 6 min Dideric Cuckoo U on severalseparate occa- Maclaren 1953 sions, different adults fed different young recently able to fly Dideric Cuckoo U c• fed fully fledgedyoung Symonsin Friedmann 1956 Dideric Cuckoo U adultsfed fledglingson sev- Smith 1957 eral occasions Dideric Cuckoo U c• fed young Thomas 1960 Dideric Cuckoo U adult fed fledgling Ottow and Duve 1965 U on severalseparate occa- Hume in Fulton 1904 sions, different adults fed different young Brown-headed Rose-breasted 9 fed nestlingdaily Bonwell 1895 Cowbird Grosbeak (k) Brown-headed Common Yellow- 9 fed cowbirdat nestling Walton 1903:219 Cowbird throat (k) and fledgling stage Brown-headed U 9 fed juvenile repeatedly Fletcher 1925 Cowbird during one day of observa- tions Brown-headed U within a flock, c• c• and 9 9 Merritt 1956 Cowbird fed many young Brown-headed Northern Cardi- and 9 cowbird and 9 car- Hernandez 1986 Cowbird nal (s) dinal fed youngcowbird

• Known host species(k), suspectedhost species(s) providedby the author, unknownhost 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 Observationsof provisioning of young by brood parasitesprovide strong evidencethat the propensityto provisionyoung has not been lost by at least some individualsof some speciesof brood parasites.This be- havior is either vestigialor a new development(Kikkawa and Dwyer 1962). If the latter, this behavior may have developed as an extension of court- ship feeding (Bensonand Serventy1957). Courtshipfeeding and "pa- rental" feeding have similar elements, although the motives for these behaviorsdiffer. Courtshipfeeding allowsfemales to assessthe food-gath- 368] J. C. Lorenzanaand S. G. Sealy J.Field Ornithol. Summer 1998

TA•I.}• 2. Parasiticspecies that have been observedprovisioning young and/or courtship feeding.

Observations of feeding Court- Parasiticspecies Young ship Courtshipfeeding references

INDICATORIDAE Greater N a N not mentioned in Friedmann 1955 d CUCULIDAE Pied Cuckoo Y Y Godfrey 1939, Liversidge1971 Y Y Channer 1976 Thick-billed Cuckoo N Y Rowan 1983 Red-chested Cuckoo N Y Rowan 1983 Black Cuckoo N Y Young 1946 Common Cuckoo Y Y Wyllie 1981 Pallid Cuckoo Y Y Klapste1981, Lord 1956, Noske 1978, Robinson 1950, White 1950 N Y Noske 1978 Fan-tailed Cuckoo Y Y Chisholm 1940, Smithers 1977, Noske 1978 ShiningBronze-cuckoo Y Y Falla et al. 1978, Serventy1958, Wat- son and Bull 1950 Horsfield's Bronze-cuckoo N Y McCulloch 1967, Moffat 1978, Wall 1978, Watson 1955 Klaas's Cuckoo Y Y Winterbottom 1939 African Emerald Cuckoo Y Y Haydock 1950 Asian Koel Y N n/a Long-tailedKoel N b N n/a Channel-billed Cuckoo Y Y Goddard and Marchant 1983

FRINGILLIDAE Bay-wingedCowbird N N not mentioned in Friedmann 1929 d ScreamingCowbird N N not mentioned in Friedmann 1929 a N N not mentioned in Friedmann 1929 a N c N not mentioned in Friedmann 1929 a Brown-headed Cowbird Y N not mentioned in Friedmann 1929 a

a Possibleobservation by Hartlaub in Friedmann (1955:162). b Possibleobservation by Cheeseman(1890). c Possibleobservation by McCracken (1984). d Courtshipfeeding was not mentioned to occur in these speciesalthough there was a sectionon courtshipbehavior in Friedmann (1929, 1955). ering ability of males, increasesthe female's nutritional reserves,and fa- cilitatespair formation (Nisbet 1973, Smith 1980). "Parental" feeding, on the other hand, contributesprimarily to the growth and development of the parasite.Courtship feeding, which alsooccurs in nonparasiticcuck- oos (e.g., Dwarf Cuckoo, Ralph 1975; Black , Rowan 1983; Black- billed Cuckoo, Spencer1943) and many other nonparasitictaxa, hasbeen reported in 11 avian orders and in almost three-quartersof families (Smith 1980). Although provisioningof young occursin many of Vol.69, No. • BroodParasites Feeding Young [ 369 the same parasitic speciesthat also courtship feed, there is not enough evidenceto suggestthat the provisioningof young developedfrom court- ship feeding. First, both male and female brood parasiteshave been ob- served provisioning young, whereas only males feed adult females. Sec- ond, the provisioningof young hasbeen observedin two parasiticspecies that do not courtship feed (Asian and Brown-headedCowbirds). Third, there is no significantassociation between courtshipfeeding and provisioningof young. It is not surprisingthat female brood parasiteshave been observed feeding nestlingsor fledglingsbecause they probably are the ones that find a nest and determine when it is suitablefor laying and, in the case of many species,visit it to remove or pierce a host (e.g., Brooker and Brooker 1989, Carter 1986, Gill 1983, Jensen and Jensen 1969, Livesey 1936, Mason 1980, Sealy1992). Males are not commonlyinvolved in any of these activities,although there are a few observationsof activity by parasitic males around nests (e.g., Sealy 1994). In light of the fact that parasiticmales are seldom observedaround host nests,it is interesting that males of the following specieshave been observedfeeding nestlings and fledglings:Klaas's Cuckoo, African Emerald Cuckoo,Dideric Cuckoo, and Brown-headed Cowbird.

PARENT-YOUNG ASSOCIATION Use of molecular genetic techniqueswould determine whether the adults are the genetic parents of the parasiticyoung they are feeding. Hahn and Fleischer(1995) found that femaleand juvenile Brown-headed Cowbirdstrapped together had higher band similaritiesbased on restric- tion fragmentsthan would be expectedby chance.This finding, however, was based on only 11 pairs of adult female and juvenile ,and individuals trapped at the same time may not have arrived at the trap together. Also, baited traps possiblyaffect the distribution of birds and may unnaturally attract many birds to a small area. Nonetheless,Hahn and Fleischer'sfinding is interesting becauseyoung cowbirdsappeared to associatewith their mothers. Fletcher (1925) observed an adult female cowbirdthat repeatedlyfed the samejuvenile although there were other young cowbirdsfeeding in the samearea. The relationshipbetween these individuals, however, was not known. Mother-offspring associationscould result from female cowbirds re- cruiting their young, or from young cowbirdsseeking out their mothers (McCracken 1984, Hahn and Fleischer 1995). The only way that young cowbirdscould become visuallyfamiliar with their mothers is if females visited the nest during the nestling stage.Female cowbirdsare known to visit nestsafter parasitismto remove host eggs(e.g., Mayfield 1961, Sealy 1992), however,mother-offspring associations cannot developwhen nests are visited during the egg stage.Young cowbirdscould become acousti- cally familiar with their mothers if the latter vocalizenear the nest (Hahn and Fleischer 1995). Females frequently are in the area of a nest they have parasitizedbecause there is evidence that they have home ranges 370] J. C. Lorenzariaand S.G. Sealy J.Field Ornithol. Summer 1998

(e.g., Alderson 1996, Darley 1983, Rothstein et al. 1984, Teather and Rob- ertson 1985). Hahn and Fleischer (1995) suggestedthat by associatingwith adult brood parasites,juveniles can better learn species-specificbehavior. How- ever, species-specificbehavior of brood parasites,such as singingin males and songrecognition in females,has been found to be innate (King and West 1977), and Friedmann (1929) reported that large flocks of Brown- headed Cowbirdsformed during the fall are frequentlycomposed entirely of juveniles. Therefore, juvenile parasiteslikely seek out individuals of their own speciesrather than being recruited by adults. There is evidence that Great Spotted Cuckoosmonitor nestsafter lay- ing. Soler et al. (1995) suggestedthat femalesof this speciesmonitor nests they have parasitizedto ensure that their egg remains in the nest. Nest , presumablyby female cuckoos,is more likely to occur if the parasitic egg has been ejected. This forces hosts to renest and provides the cuckooswith another chanceto lay an egg (see also Zahavi 1979). Similarly, Arcese et al. (1996) suggestedthat cowbirdsdepredate nests that are discoveredtoo late in the host'snesting cycle for parasitismto 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 displayat the approach of a potential predator to a parasitizednest, and Gabrielson (1921) described another instance of this behavior. Fe- males should benefit by monitoring the progressof a nest and protecting the nest from the approach of predators, but the costsof spending so much time in this activity,however, likely outweigh the benefits. The scar- city of reports of nest protection by female cowbirds suggeststhat this behavior is rare. The female cowbirdsprobably 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 Cuckoosappears to be more common. Soler et al. (unpubl. data in Soler et al. 1995) were scold- ed by cuckooson 25% of 56 visitsto 30 parasitizedBlack-billed Magpie nests,whereas they were scoldedby magpieson only 5% of the visits.

STRATEGY OR OPPORTUNISM. > About 90% of the recordsof provisioningin brood parasitesinvolved cuckoos.This may reflect the fact that courtship feeding is found only among the cuckoos.Evidence for this, however,is weak becausewe did not find a significant associationbetween courtship feeding and provi- sioning behavior.More likely, it is becausethere are more speciesof par- asitic cuckoosthan cowbirdsand . The question remains whether the provisioningof young by brood par- asitesis common enough to be considered a strategyor whether it is simply trivial behavior.Like other speciesof birds, brood parasitesmay possessan innate responseto feed begging young (see Craig and Jamie- son 1990). If parasiticadults respond to the stimulusof begging young, Vol.69, No. 3 BroodParasites Feeding Young [371 then there should alsobe observationsof adult parasitesfeeding begging young of other species.We did not find observationsof this behavior, although there are many casesreported of nonparasiticspecies feeding other nonparasitic species (Shy 1982), and individuals other than the original foster parents of parasiticspecies feeding parasiticyoung (Sealy and Lorenzana1997). The lack of suchobservations suggests that it is the adult brood parasite'sintention to feed an individual of its own species. Implicitly, the adults recognizeyoung of their own species. Provisioningof young by brood parasitesmay be important if hosts cannot provide the young parasitewith enough or the right kind of food. In such cases,limited parental care by brood parasitescould be an ad- aptationto poor-qualityhosts. Benson and Serventy(1957) suggestedthat provisioningby brood parasitesmay be necessaryin caseswhere insectiv- orous brood parasiteslay eggsin nestsof herbivorousspecies. There is, however,no evidence for this becauseparasitic nestlings raised by herbiv- orous speciesusually do not surviveto fledging (Eastzeret al. 1980, Kozlo- vic et al. 1996, Middleton 1991; but see Seel and Davis 1981). Further- more, most observationsinvolve brood parasitesfeeding fledglings,not nestlings.Brood parasiticspecies should be stronglyselected to avoidher- bivoroushosts altogether (Kozlovicet al. 1996), rather than to compen- satefor herbivoroushosts by feeding the young at the nest.

ACKNOWLEDGMENTS

This review was initiated during a sabbaticalleave granted to SGS by the University of Manitoba. N. L. Sealyprovided invaluablehelp with the early stagesof the literature search. Commentsfrom J. V. Briskieand C. R. Chandler greatlyimproved the manuscript.Financial support was provided by a NSERC research grant to SGS and NSERC and (Universityof Manitoba) Facultyof ScienceUndergraduate Summer Research awards to JCL.

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APPENDIX. Common and scientificnames of avianspecies mentioned in the paper.a

FAMILY Common name Scientific name

INDICATORIDAE indicator CUCULIDAE Pied Cuckoo Oxylophusjacobinus Great Spotted Cuckoo Clamatorglandarius Thick-billed Cuckoo Pachycoccyxaudeberti Red-chested Cuckoo solitarius Common Cuckoo C•t. canor,rs Pallid Cuckoo Cu. pallidus Brush Cuckoo variolosus Fan-tailed Cuckoo Ca. flabelliformis Shining Bronze-cuckoo Chrysococcyxlucidus Klaas's Cuckoo Ch. klaas African Emerald Cuckoo Ch. cupreus Dideric Cuckoo Ch. caprius Asian Koel Eudynamysscolopacea Long-tailed Koel E. taitensis Channel-billed Cuckoo Scythropsnovaehollandiae CENTROPODIDAE Black Cuckoo Centropusgrillii COCCYZIDAE Dwarf Cuckoo Coccyzuspumilus Black-billed Cuckoo Co. erythropthalmus CORX•qDAE Black-billedMagpie Pica pica MELIPHAGIDAE White-plumed Lichenostomuspenicillata PARDALOTIDAE White-browed Scrubwren Sericornisfrontalis Striated Calamanthus Calamanthusfuliginosus Yellow-rumpedThornbill Acanthiza chrysorrhoa Grey Gerygone Gerygoneigata VIREONIDAE Red-eyedVireo Vireo olivaceus PASSERIDAE House Sparrow Passer domesticus Meadow pipit Artthuspratensis Village Weaver Ploceus cucullatus FRINGILLIDAE Yellow Warbler Dendroicapetechia Common Yellowthroat Geothlypistrichas Rose-breasted Grosbeak Pheucticus ludovicianus Northern Cardinal Cardinalis cardinalis Bay-wingedCowbird Molothrus badius ScreamingCowbird M. rufoaxillaris Shiny Cowbird M. bonariensis Bronzed Cowbird M. aeneus Brown-headed Cowbird M. ater

Order and nomenclaturefollow Sibleyand Monroe (1990).