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j. Field Ornithol., 53(4):370-393

INTERSPECIFIC FEEDING AMONG : A REVIEW

BY MARILYN MUSZALSKI SHY

Much recentattention has been devotedto intraspecifichelpers at the nest. Skutch(1935, 1961, 1976) listed severalinstances of interspecific helping, but a thoroughconsideration of this topic is lacking.The pur- poseof this paper is to give a comprehensivesurvey of the occurrence of the feeding of one by another of a different speciesand to describe the conditions under which the behavior has tended to occur. The subjectis potentiallyof considerableevolutionary interest since in interspecifichelping, kin selectionis impossible. Certain instances have been omitted from the discussion and the list. They include captiveand parasiticbirds and situationswhere a clutch has been partially or fully replacedwith that of another speciesby an experimenter.Common names for bird specieswill be usedthroughout the paper; a list of scientificnames appears in Appendix I.

RESULTS

The list of instancesof interspecificfeeding shownin Table 1 was compiledby searchingthe literature (Appendix II) and requestsmade through severalornithological journals. Table 1 summarizes140 cases of interspecificfeeding. Adopting species are representedby 22 families and 65 species,and birds that were fed by 22 familiesand 71 species (Tables2, 3). There were 95 casesof nestlingsfed, 30 casesof fledglings, 11 casesof both nestlingsand fledglings,and 4 caseswhere no age was given. I have classifiedinstances of interspecificfeeding into 8 categories related to their probableproximate causes(Table 2): (1) for somerea- son,the bird wasraising a mixed clutch;(2) the original nestand brood of the bird were destroyed;(3) the nestof anotherspecies was very close to that of the bird performing the behavior; (4) young birds calling stimulatedanother species to feed them; (5) orphanedbirds were adopt- ed temporarilyor permanently;(6) a male bird fed another specieswhile his mate incubated;(7) finding a matelessbird, or beingmateless itself, a bird joined a heterospecificindividual or pair with young; (8) a mis- cellaneouscategory: none of the abovereasons were evident. Not all of the categoriesare mutuallyexclusive. Mixed clutches.--Mixedclutches have been frequently observed,es- peciallyamong hole-nestingspecies when competitionfor nestingsites is severe.Mackenzie (1954) deliberatelyremoved half of 60 boxesin an area during the fall in an attempt to study effects of competitionon cavity-nesters.The followingspring, a Redstartpresumably laid eggsin 2 tits' nests.The Great and Coal Tit parentsraised the young Red- startsalong with their own. Great Tits havebeen thoughtto usurp Blue 370 Vol.•3, •o. 4 InterspecificFeeding [371

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TABLE 2. Incidenceof proximatereasons for interspecificfeeding.

Reason No. of cases Percent a

Mixed clutch 32 21.2 Originalnest destroyed 7 4.6 Closenest of another species 36 23.8 Callingby young 15 9.9 Orphaned birds 6 4.0 Mateless birds 7 4.6 Male, mate incubating 8 5.3 Miscellaneous 40 26.5

a Percentagesreflect reasons cited; in somecases more than one reasonwas given.

Tit nests(Arn 1955, Weinzierl 1958). Occasionally,Great Tits have laid in other nests,including those of the Blue Tit and (Arn 1955). Amann (1949) found a number of similar casesin previous years. Gustavsson(in Mackenzie 1954) found a banded Coal Tit and banded Marsh Tit had laid in the samenest; the Marsh Tits ended up raising a mixed clutch of 4 Marsh Tits and 2 Coal Tits. It was unclear whether the Marsh Tits oustedthe originallynesting Coal Tits, or the Coal Tits deserted or were killed and the Marsh Tits took over the nest. At least3 casesof AmericanRobins sharing a nestwith anotherspecies have been observed:with a Mourning Dove (Raney 1939); with a Gray Catbird (Benton 1961); and with a House (Bailey and Niedrach 1936). A Robin shared a nest with a Pied (Lack 1953), and a Cardinal and SongSparrow raised a combinedbrood (Brackbill 1952). Nestor brooddestroyed.--There are severalinstances of birds feeding another speciesafter their own nest was destroyed.In one, a pair of Robins fed Song Thrushes (Lack 1953). Southern (1952) observeda pair of SpottedFlycatchers feeding Blackbirdsafter their nest wasde- stroyedin a storm.After a pair of Starlingsrepeatedly lost a nestplaced on a drainspout,they fed nearby nestlingAmerican Robins (Putnam 1961). A male Cardinal fed American Robins after his first nest was destroyedand whilehis mateincubated their secondclutch (Logan 1951). Song Sparrowsfed Yellow Warblers while nearby there was a Song Sparrownest with dead young in it, presumablybelonging to this pair (•lackson1941). When a researchertook the eggsof a ScreechOwl, the brooded flickersin a hole in the same tree, and even brought a small bird for the nestlingsto eat (Lyon 1922). A Starlingfed nestlingPurple Martins after 4 eggswere taken from the martin-houseand destroyed (Brown 1977a). This categoryis probablylarger than it appears.In many observations,summarized later, the nestinghistory of the birds performing the behaviorwas unknown. Closenesting of anotherspecies.--There are many examplesof 2 species nestingclose to each other and one or both taking an interest in the nest of the other. In some cases,a pair has attackedanother species Vol.53, No. 4 InterspecificFeeding [38 1 bringing food to their nestlings;at other times the pairs took turns feedingall youngwith no antagonism. It is not known how frequentlya nest might fail due to one or both parents neglectingit for another nest. A male Pygmy Nuthatch fed nestlingMountain Bluebirdsin a nest .5 m abovehis own; his own nest failedas he wasattracted to the bluebirdscalling before his hatched (Pinkowskipets. comm.).A male EasternBluebird fed nestlingHouse Wrensas his mate incubated and later neglectedhis ownyoung; he even fought the parentsto feed the youngbirds (Forbush1929). A pair of Arctic Loonsraised a brood of 5 SpectacledEiders (Abra- ham 1978). Initial observationsincluded the locatingof the and ducknests 10 m apart. One monthlater, the duckswere being fed by the ,although Spectacled Eiders are self-sufficientupon hatching. The fate of the originalloon nestwas not known. The timing of the hatchingof the 2 nestsseems to be a criticalfactor. If 2 pairsof birdsnest close to eachother, the nestlingsthat hatchfirst may receivethe attentionof one or both of the birds of the other pair. This was the casewhen a male Tree Swallowwas observedfeeding young American Robinsin a nest atop the housein which his own nest was located(Munro 1929). The nestlingswallows were about 1 week youngerthan the robins.Males will sometimesfeed at anothernest while their own nestis beingincubated. A HouseWren fed nestlingCommon Flickers while his mate incubated and continued to do so after his own eggshatched (Royall and Pillmore 1968) and a Wren fed Great Tits under similar circumstances(Armstrong 1955). This will be discussed further in a separatecategory. In one sense, the fact that one nest is close to another cannot be considereda reasonfor the interspecificfeeding behavior since in any givenarea, there will alwaysbe a numberof speciesnesting close to each other.Only thoseinstances are citedwhere the originalauthor believed it significantenough to mention. Its importanceis due to the fact that it indicatesthat the bird is breeding;in somecases individuals perform- ing the behaviormay not be. Youngbirds calling as a stimulus.--Incertain cases of interspecificfeed- ing, the observersbelieved the callsof the young were an important factor triggeringthe behavior.In somecases, fledglings or nestlings were fed as anotherbird wasforaging nearby. Fledglings have directly solicitedfood from adults of a different species.A Tufted Titmouse beggedand ran towarda Red-belliedWoodpecker, which fed it (Curry 1969). While a male Hairy Woodpeckerwas feeding at a suet holder, an immature Downy Woodpeckercalled for food and was fed by the Hairy (Davis 1973).A fledglingPied Wagtailchased a Blackbirdand the Blackbirdfed it 6-7 times(Russell 1973). A Worm-eatingWarbler fed a fledglingKentucky Warbler as it calledfrom the ground(Hickey pets. comm.). Of course,when a nest of one speciesis near the nest of a second(as in the precedingcategory), the visualand auditorysignals could togeth- er triggerfood-bringing behavior. Begging by nestlingsmust rarely elicit 382] M.M. Shy j. F•eldOrnithol. Autumn 1982 feedingby heterospecificbirds, based on the scarcityof suchobservations in the manypublished studies of nestingbehavior. Studies of the feeding of fledglingsare muchrarer, soit is uncertainwhether feeding of fledged young is a rare event. It is not likely that unsuccessfulpanhandling attemptswill be reported in the literature; however, they do occur. Brewer (pers.comm.) saw a fledglingYellow Warbler beg from a female , which peckedat it. Orphanedbirds.--In severalcases, one specieshas brought food to or- phaned broods or individualsof another species.This could be trig- geredby the presumablyloud callingof unsatiatedyoungsters. Snyder (1913) noticeda broodof EasternKingbirds calling loudly after an elec- trical storm. The parents were not seen afterward. An Eastern Wood Peweefed the orphansfor 10 days,until they fledged.A male Robin fed motherlessBlackbirds while his mate was incubating,after which the male Blackbirddeserted the nest.The Blackbirdsfledged under the Robin'scare (Armstrongand Whitehouse1977). Black-and-whiteWar- blers fed an fledgling which had becomeseparated from its parents (Kendeigh 1945). A pair of Chipping Sparrowsfed a brood of fledgling Purple Fincheswhile continuingto feed their own brood in a nestin a smallspruce one tier of branchesbelow the finch nest(Jackson 1941). Malefeeding another species while mate incubates.--Two previous sections includeseveral cases of malesfeeding nestlings of anotherspecies while their mateswere incubating.Wight (1934) noted a CarolinaWren feed- ing Great Crested Flycatchersfor a short time, although after a few fights with the parents,the wren gave up the attempt. A male Scarlet fed Chipping Sparrowsbefore his own eggs hatched (Hales 1896), and a Dark-eyedJunco fed Bewick'sWrens ashis mate incubated (Lonsdale 1935). Skutch(1961) noted certainmales are soeager to begin feeding their nestlingsthat they offer food to the unhatchedeggs. He suggestedthat feeding of another species"may provide an outlet for repressedener- gy." Matelessbirds.--A few casesof interspecificfeeding have been docu- mentedthat involveda bird that had beenunsuccessful in finding a mate, or had joined a parent bird whose mate had disappeared.A female Northern Three-toed ,out of the known breeding range of this species,fed nestlingBlack-backed Three-toed while they called loudly from the nest (Hickey pers. comm.). After several days of observation,Possert (1955) concludedthat only 2 birds were attendinga Great Tits' nest,a male Great Tit and a Long-tailedTit. A male House Wren that could not securea mate, althoughhe had filled a birdhousewith nestingmaterial, fed 3 Black-headedGrosbeaks until theyfledged, and then fed nestlingHouse Sparrows (Hills 1924).A male Blackbirdfed 2 nestlingSong Thrushes,assisting the female ; the nestlingsfledged and were observedfor severaldays with the Black- bird (Moore 1973). A banded male Dusky SeasideSparrow that had bred successfullyin previousyears, could not locatea matedue to a lack Vol.53, No. 4 InterspecificFeeding [383 of females (Rakestrawpers. comm). (At that time there was only one female in the entire area of St. John's National Wildlife Refuge near Titusville,Florida.) It fed 2 fledglingRed-winged Blackbirds for 4 days. A male EasternBluebird joined a Mountain who had losther mateand helpedher raiseher brood(Scott 1971). A Worm-eatingWar- bler fed nestlingOvenbirds at a nest where only one parent Ovenbird was observed (Maciula 1960). Miscellaneouscategory.--This is a large category(Table 2) but may only appear so due to omissionsin either observationsor recordsof observ- ers. There are, nevertheless,some unusual circumstancessurrounding certain instancesof interspecificfeeding in this category. Under the categoryof mixed clutches,it was mentioned that some birds are thought to usurp nests,in somecases with a partial clutch, especiallyin casesof severecompetition for nest sites.Hovingh and Ponshair(1951) cited this as a possiblereason for a Great Horned Owl raisinga brood of 3 Red-tailedHawks. A female CommonFlicker may haveusurped a Starlingnest, as shewas observed caring for 3 nestling Starlingsin what appearedto be a typical Starling nest (Lott 1939). A Mourning Dove, whoseown eggsfailed to hatch, began to care for a brood of White-wingedDoves, who had been neglectedby their own parents (Neff 1945). Perhapsone of the most unusual casesis that of a female Blackbird that, after rearing her own young,continued to offer food for 2-3 weeks to any bird comingnear. A Robin acceptedthe food (Lack 1953). Taxonomicand ecologicaldistribution. roOf the 65 speciesthat were ob- servedfeeding another species, 25 havemultiple records(Table 3). The highest number of observationsfor a specieswas 10, for the House Sparrow,although it wasobserved only twice to be fed by anotherspecies. Of speciesthat were fed, there are multiple recordsfor 29. Most fre- quently observedwas the , with 11 records. The 22 families of birds fed are listed in Table 4. There are 4 instancesof one speciesfeeding another, with the reverse situationalso being observed.These speciespairs are (excludingcases of mixed clutches)Common Flicker--Starling, Purple Martin--Starling, Robin--Blackbird, and American Robin--House Finch. Comparisonsby habitat,diet, and type of nest,indicate similarities between"adopters" and speciesbeing fed (Table 5). Edge specieswere mostoften observed.This is likely due to the nature of the observations, many made by suburbanbackyard birdwatchers. Of 8 casesof warblers feedinganother species,5 were ground-nesters.Perhaps the list of ob- servationsreflects birds that are most easilyseen, and implies the pos- sibilityof other, not so visiblespecies, being just as likely to engagein the behavior.

DISCUSSION The interspecificfeeding observedas a result of mixed clutchescan- not be consideredin the sameway as under the other conditions.Peek et al. (1972) have shownthat Red-wingedBlackbirds do not appearto 384] J. Field Ornithol. Autumn 1982

TABLE3. Specieswith multiple records.

No. of cases

Species Adoptors Birds fed

Red-tailed Hawk 0 2 Mourning Dove 2 0 Common Flicker 0 4 EasternKingbird 0 3 Tree 2 2 Purple Martin 2 0 Pied Wagtail 0 2 2 0 Winter Wren, Wren 7 4 House Wren 2 4 Gray Catbird 4 3 0 2 Robin 7 4 Redstart 2 4 4 1 0 2 Blackbird 3 5 Song Thrush 0 2 American Robin 6 11 SpottedFlycatcher 0 2 Marsh Tit 2 0 Mountain Chickadee 2 0 Coal Tit 0 2 Great Tit 6 5 Blue Tit 7 7 Tufted Titmouse 0 3 Nuthatch 3 0 Song Sparrow 4 0 Dark-eyedJunco 0 2 Chipping Sparrow 0 3 Field Sparrow 0 3 Rufous-sided Towhee 4 0 Brown Towhee 2 0 Cardinal 4 3 Black-and-white Warbler 2 0 Yellow Warbler 0 2 Ovenbird 0 2 Worm-eatingWarbler 2 0 House Finch 0 3 House Sparrow 10 2 Starling 4 4 Total number of specieswith multiple records 25 29 recognizetheir own youngeither in or out of the nestuntil they are 10 daysold. In 2 of 3 albatrossspecies, parents did not appearto recognizea strangechick put into their neststo replace one of their own chicks (Tickell and Pindar 1972).Since the chicksrarely leavetheir own nests, there seemsto be no advantagein learningto recognizea bird's own Vol.53, No. 4 InterspecificFeeding [385

T^BI•E4. Familiesof adoptorsand birds fed, and number of speciesfor each.

No. of adopting No. of species Family species of birds fed

Gaviidae 1 0 0 1 1 1 1 0 1 2 Strigidae 2 1 Trochilidae 1 1 Picidae 4 4 Tyrannidae 5 2 Hirundinidae 2 3 2 1 Troglodytidae 3 3 Mimidae 1 4 Prunellidae 0 1 Muscicapidae 9 13 1 0 Paridae 7 6 Certhiidae 0 1 Sittidae 2 0 Emberizidae 11 9 Parulidae 6 7 Vireonidae 0 1 Icteridae 1 3 Fringillidae 2 5 1 1 Sturnidae 1 1 Total numberof species 65 71 young until later, perhaps upon fledging. This idea is supported in a study of factors relating to the timing of parent-chick recognitionin (Burtt 1977).Birds may feed almostanything that happensto be in their nestsunder the right conditions.It is of interestthat birds fed and, in somecases, successfully reared another speciesin their own nests,but it doesnot explain spontaneousinterspecific feeding. Becausethere are so many casesof what seemsto be an unusualand not easilyobserved phenomenon, especially for fledglings,it would ap- pear the behaviorrequires an evolutionaryexplanation. One possibility is that the behavioris non-adaptive,and a consequenceof other adaptive features.For example,it may be ordinarilyadvantageous to have such a strongdrive to care for young that it overcomesheavy deterrents. If this is so, feeding inappropriateyoung may be rare enough,and carry a smallenough penalty, that no evolutionarymodification of the strong drive occurs. Arguing againstthis possibilityis the fact that it is difficult to provoke the feeding of alien youngin any fashionexcept by placingthem in the parentalnest. When parent swallowsbegin actively to discriminatetheir 386] M.M. Shy j. FieldOrnithol. Autumn 1982

T^BLE 5. Number of speciesof adoptorsand birds fed, accordingto habitat, diet, and type of nest.

No. of adoptingspecies No. of speciesof birds fed

Habitat Edge 41 50 Fores t 19 17 Water 3 2 Grass 2 1 Desert 0 1 Primary Diet Insectivorous 45 51 Granivorous 13 16 Other 7 4 Type of Nest Open 39 45 Cavity 26 26

own young from others(upon fiedging)they are aggressiveto alien chicks (Burtt 1977). Dawkins (1976) suggestedthat adoption should be a rare mistake, sinceit confersno benefitsupon the foster parents,but insteadwastes time and energy that could be investedin their own kin. He did, how- ever, allow that adoptorscould benefit by gaining practicein the "art of childrearing."From this hypothesis, one would predict younger birds to be more likely to engagein interspecificfeeding than older birds, sincethe experiencethey would gain by the feeding practicewould directlybenefit them and enablethem to be better parents. That experiencedoes improve nesting success is suggestedby a study conductedby DeStevens(1978). She found that youngerfemale Tree Swallowshave lower fiedging rates for their broodsthan older female Tree Swallows.Lehrman and Wortis (1967) presentedconclusive evi- denceof experienceimproving parental care in RingedTurtle Doves, anda numberof similarobservations support this idea as well (Hediger 1950, 1955, Lehrman 1961). In mostcases of interspecificfeeding, the age of the helper is, unfor- tunately,unknown. Porcher(pers. comm.)mentioned a caseof a juve- nile HouseSparrow feeding nestling Tree Swallows.He observedolder sparrowsbeing driven off by swallows,and believedthe young sparrow wasallowed to approachthe nest becauseit wasnot yet mature. Several casesreport definitelyolder, experiencedbirds feeding another species. Rakestraw's(pers. comm.) observationof the Dusky SeasideSparrow that had bred successfullyin previousyears is a casein point. The bird had had ample experiencein raisingyoung. The male Blackbirdwho assisteda matelessSong Thrush was"mature" (Moore 1973). It may be more advantageousfor a youngbird to feed anotherspecies Vol.33, Xo. 4 InterspecificFeeding [387

than not to engagein the feeding behaviorat all if experienceimproves an organism'sparental abilities. In certaincases, this might be especially true, for example, when a bird's own nest failed and it wastoo late for it to renest,or when it's own mate had been lost and it could not readily find another, or when a male bird, possiblyinexperienced, has time to practiceon its neighbors'offspring while his mate incubates.Other hy- pothesesexplaining interspecific feeding are conceivable,such as recip- rocalaltruism (Trivets 1971), but the currentlyavailable information seemsnot to provideany ready way of approachingthem. Although acceptingfood no matter who the donor is should be ad- vantageousin most circumstances,being raised by foster parentsmay lead to later problems.When a pair of GommonTerns raiseda Herring , it did not behaveas a normal gull (Kuhlemann 1939). A Mourning Dove remained with its foster family of Ringed Turtle Doves rather than joining its own species(Grewe 1959). Female Zebra Finchesraised by BengaleseFinches showed no clear preference for their own species in choosinga mate(Sonnemann and Sj/51ander1977). Herring and Less- er Black-backedgulls showedabnormal migration and hybridization after cross-fosteringexperiments (Harris 1970).These types of reactions indicatethat cross-fosteringmay not be totallybeneficial to the adopted bird.

ACKNOWLEDGMENTS I thank the followingwho contributedobservations of interspecific feedingbehavior cited in this paper: Ken Gammon,Joseph J. Hickey, D. Edward Holman, Ben Pinkowski,Alfred G. Porcher,and JamesL. Rakestraw.I am alsovery gratefulto RichardBrewer who providedthe impetus for the paper and also aided in severalrevisions of the manu- script,and to RichardPippen and JamesErickson whose suggestions were most helpful.

LITERATURE CITED

ABRAHAM,K.F. 1978. Adoptionof SpectacledEider ducklingsby ArcticLoons. Condor 80:339-340. AMANN,F. 1949. Junge Kohl- und Blaumeisenim gleichenNest. Ornithol. Beob. 46: 187-190. AMERICANORNITHOLOGISTS' UNION. 1957. 5th ed. Lord Baltimore Press, Baltimore, Maryland. ANTEVS,A. 1947. Towheehelps Cardinals feed their fledglings.Condor 49:209. ANTOINE,N.J. 1959. Blue Tit feeding young .Br. Birds 52:432-433. (abstractin Bird-Banding31:100). AOU CHECKLISTOF NORTH AMERICANBIRDS. 1957. 5th ed. Lord Baltimore Press, Bal- timore. ARMSTRONC,E.A. 1955. The Wren. Collins, London. --, ANDH. L. K. WHITEHOUSE.1977. Behaviouraladaptations of the Wren (Trog- lodytestroglodytes). Biol. Rev. 52:235-294. ARN, H. 1955. Mischbruten von Kohlmeisen, Blaumeisen und kleiber. Ornithol. Beob. 52:129. (abstractin Bird-Banding28:106). AUSTIN,O. L. 1958. Life historiesof North American cardinals,, buntings, towhees,, sparrows and their allies; Passeriformes,family Fringillidae. U.S. Natl. Mus. Bull. 237. 388] M.M. Shy J FieldOrnithol. Autumn 1982

BAILEY,A.M., ANnR. j. NtEr•mac•. 1936. Communitynesting of WesternRobins and House Finches. Condor 38:214. BAILLAmCE,W. 1930. A BronzedGrackle foster parent. Can. Field-Natß44:166-167. BATTS,H. L., JR. 1958. The distributionand populationof nestingbirds on a farm in southern Michigan.Jack-Pine Warbler 36:131-149. BEca,H. H. 1925. ProthonotaryWarbler feedingyoung of anotherspecies. 42: 140. BENTON,A.H. 1961. Nest sharingby robin and catbird. Kingbird 11:81-82ß BETTS, M. M. 1958. The behavior of adult tits toward other birds and mammals near the nest. Br. Birds 51:426-429. BRaCaBILL,H. 1952. A joint nestingof Cardinalsand SongSparrows. Auk 69:302-307. BRace, M.B. 1968. Kingbird feedingBaltimore Oriole nestlings.Auk 85:321. BRINDLEY,M.D. 1937. House Sparrowfeeding young Hedge Sparrows.Br. Birds 31: 25. BRooas, E. 1922. Cardinal and catbird. Bird-Lore 24:343-344. BROWN,C.R. 1977a. Starling'feedingPurple Martins.Southwest. Nat. 21:557-558ß ß 1977b. Unusual.feedingbehavior of a male Purple Martin. Bull. Tex. Ornithol. Soc. 10:15-16. BURtESS,D.D. 1964. LeastFlycatcher "assists" at Chipping Sparrownest. Kingbird 14: 216. BURTT,E. H. 1977. Some factorsin the time of parent-chickrecognition in swallows. Anim. Behav. 25:231-239. CARR,T., ANn C. j. GoiN,JR. 1965. Bluebirdsfeeding Mockingbirdnestlings. Wilson Bull. 77:405-407. C•APMAN,L.B. 1955. Studiesof a Tree Swallowcolony (third paper). Bird-Banding 26:45-70. CURRY,J. R. 1969. Red-belliedWoodpecker feeds Tufted Titmouse. WilsonBull. 81: 470. DAVIS,G. L. 1973. Hairy Woodpeckerfeeding immature Downy WoodpeckerßKing- bird 23:189. DAWaINS,R. 1976ß The selfishgene. Oxford UnivßPress, New York. DECa,R.S. 1945. The neighbor'schildren. Nature Mag. 38:241-242. DESTtVENS,D. 1978. The influenceof age on the breedingbiology of the Tree Swallow Iridoprocnebicolor. 20:516-523. FITCH,H.S. 1949. Sparrowadopts kingbird. Auk 66:368-369. FORBUS•,E. H. 1929. Birds of Massachusettsand other New England states.Vol. 3. Mass.Dep. Agric., Boston. GREENLAW,J. S. 1977. White-throatedSparrow as fosterparent of fledglingDark-eyed Junco.Bird-Banding 48:170-171. GREWE,A. 1959. Ring Dovesrear Mourning DoveßFlicker 31:24. HALES,H. 1896ß Peculiartraits of someScarlet . Auk 13:261-263. HAMILTON,G.D. 1952. EnglishSparrow feeding young Eastern Kingbirds. Condor 54: 316. HARRIS,M.P. 1970. Abnormalimmigration and hybridizationof Larusargentatus and L. fuscusafter interspecificfostering experiments. Ibis 112:488-498. HAYWARn,W.J. 1937. Incidentsin bird behavior.Wilson Bull. 49:47. HtI)ICtR, H. 1950ß Wild animalsin captivity.Butterworth, Londonß ß 1955. Studiesof the psychologyand behaviourof captiveanimals in zoosand circuses. Butterworth, London. HENr•ERSON,J. 1925. Robin feedsyoung House Finch. Auk 42:144. H•R•ERT, K. G.S. 1971. Starlingfeeds young robins. Wilson Bull. 83:316-317. HILLS,V.G. 1924. A House Wren adoptsa family of young Black-headedGrosbeaksß Auk 41:615-616. HOE•L, O. 1940. Haussperlingals Girlitzamme.Beitr•ige zur Fort-pflanzungsbiologie der Vogel 16:64-65. (abstractin Bird-Banding19:177). HOVINCI4,P. ANnJ. PONS•AIR.1951. Letter to the editor.Jack-Pine Warbler 29:88. HOYT,J. S. 1948. Observationson nestingassociatesß Auk 65:188-196. JACKSON,R. E. 1941. SongSparrows assume role of fosterparents. Bull. Mass.Aud. Soc. 25:134-135. Vol.53, No. 4 InterspecificFeeding [389

JENSEN,J. K. 1925. Mountain Chickadeewith an adopted family. Auk 42:593. JEWETT,S.G. 1928. Assistantparentage among birds. Condor 30:127-128. KENDEIGH,S.C. 1945. Nestingbehavior of wood warblers.Wilson Bull. 57:145-164. KUHLEMANN,P. 1939. Beobechtungenan einer durch Flusseeschwalben(Sterna hirundo) ausvertauschtem Ei erbr6tetenund aufgezogenSilbermove (Larus argentatus). Z. Tier- psychol.3:75-84. LACK,D. 1953. The life of the Robin. Books, London. LAWRENCE,L. I•E K. 1948. A whitethroattrio and a warbler incident.Bird-Banding 19: 122-123. LEHRMAN,D. S. 1961. Hormonal regulation of parental behavior in birds and infra- human mammals.Pp. 1268-1382, in Sex and Internal Secretion,W. C. Young, ed. 1268-1382. Williams & Wilkins, Baltimore. , ANI•R. P. WORTIS.1967. Breedingexperience and breedingefficiency in the Ring Dove. Anim. Behav. 15:223-228. LOCAN,S. 1951. Cardinal,Richmondena cardinalis, assists in feedingrobins. Auk 68:516- 517. LONSI•ALE,W.S. 1935. Blue Tits feedingyoung Robins. Br. Birds 29:113-114. LOTT,W. 1939. A flicker with a family of .Bird-Banding 10:90. LOWTHEmP.E. 1975. Scissor-tailedFlycatcher feeding Common Grackle young. Kans. Ornithol. Soc. Bull. 26:7. LYON,W.I. 1922. Owl kidnapsyoung flickers. Wilson Bull. 34:230-231. MACIULA,S.J. 1960. Worm-eatingWarbler "adopts"Ovenbird nestlings. Auk 77:220. MACKENZIE,J. m.D. 1954. Redstartsreared in tits' nests.Scot. Nat. 66:146-164. MARTIN,J. 1968. Karoo Priniasfeeding Layard's Tit-babbler. 39:263. MooKE,M. 1973. Male Blackbirds(Turdus merula) helping to rear youngSong Thrushes (T. philomelos)Br. Birds 66:365. MORON¾,J. j., w. J. BocK, ANI•j. FARR^NI•,JR. 1975. Referencelist of birds of the world. Am. Mus. Nat. Hist., New York. MOUNTFORT,G. 1957. The Hawfinch. Collins, London. MUNRO,J.A. 1929. Male Tree Swallowfeeding nestling robins. Condor 31:75-76. MURPHY,F. 1968. Chickadeeshatch and raisea bluebird. Migrant 39:11-12. NEFF,j.A. 1945. Fosterparentage of a MourningDove in the wild. Condor47:39-40. NOLAN,V., JR. 1965. A maleCardinal helper at a nestof Yellow-breastedChats. Wilson Bull. 77:196. --, ANI•R. SC•NmI•ER. 1962. A catbird helper at a House Wren nest.Wilson Bull. 74:183-184. OLENI•ORFF,R.R. 1974. Interspecificfeeding of fledglingHouse Finchesby adult Ru- fous-sided Towhees. Murrelet 55:9. PARKER,R. 1973. Sparrowfeeds young Blackbirds. Bird Life (Oct.-Dec.):31. PEEK,F. W., E. FR•N•S, ANI• D. CASE. 1972. Recognitionof nest, eggs,nest site, and youngin female Red-wingedBlackbirds. Wilson Bull. 84:243-249. PETERS,J. L. 1931. Check-listof birds of the world. Vol. I. Harvard Univ. Press,Cam- bridge. PIKE, O.G. 1930. Rambles in Britain's bird-land. London. ß 1932. The Nightingale:its storyand song.Arrowsmith, London. POSSERT,A. 1955. Schwanzmeise nilft bei der Aufzucht einer Kohlmeisenbrut. Ornithol. Beob.52:96. (abstractin Bird-Banding28:105). POWELL,F. 1927. HouseSparrow feeding nestling Spotted Flycatchers. Br. Birds21:61. POWELL,H. 1946. Nuthatchfeeds nestling Starlings. Br. Birds 39:316. (abstractin Ibis 89:152). PRESCOTT,K.W. 1967. Unusualactivities of a HouseSparrow and a BlueJayat a Tufted Titmouse nest. Wilson Bull. 79:346-347. 1971. Unusualactivity of Starlingsat a Yellow-shaftedFlicker nest. Wilson Bull. 83':195-196. PULLMAN,J. O. 1970. A Tufted Titmouse nestattended by CarolinaChickadees. Chat 34:22. PUTNAM,W.L. 1961. Starlingfeeds nestling robins. Can. Field-Nat. 75:52-53. 390] M.i. Shy J.Field Orn,thol. Autunm 1989

RANEY,E.C. 1939. Robin and Mourning Dove use samenestß Auk 56:337-338ß RE^,G. 1945ß Black-and-whiteWarbler feedingyoung of Worm-eatingWarblerß Wilson Bull. 57:262ß ROBINSON,G.G. 1962ß Winter Wren feedsTownsend's Solitaire youngß Condor 64:240ß ROYALL,W. C., JR., ANDR. E. PILLMORE.1968. House Wren feeds Red-shaftedFlicker nestlings.Murrelet 49:4-6. RUSSELL,M. 1973. Blackbirdfeeds wagtail. Bird Life (Oct.-Dec.):31. RUSSELL,W. C. 1947. Mountain Chickadeefeeding young Williamson'sSapsuckers. Condor 49:83. SAUNDERS,A.S. 1918. Blue-wingedWarbler feedinga youngField Sparrow.Auk 36: 291. SCOTT,L. 1971. Male EasternBluebird assistsfemale Mountain Bluebirdin raising young. Blue 29:126-127. SKUTCH,A. F. 1935. Helpersat the nest.Auk 52:257-273. 1960. Life histories of Central American birds. II. Pac. Coast Avif. No. 34:1- 59'3. 1961. Helpers among birds. Condor 63:198-226. ß 1976. Parentbirds and their young.Univ. of Texas Press,Austin. SNYDER,W. E. 1913. The wood peweeas a foster parent. Auk 30:273. SONNEMANN,P., ANDS. SJOLANDER.1977. Effectsof cross-fosteringon the sexualim- printing of the femaleZebra Finch (Taeniopygia guttara). Z. Tierpsychol.45:337-348. (abstractin Bird-Banding49:289). SOUTHERN,J. 1952. SpottedFlycatchers feeding nestling Blackbirds. Br. Birds45:366. SUMNER,F.A. 1933. Young SparrowHawks and a ScreechOwl in the samenest. Condor 35:231-232. SUTTON,G. M. 1968ß Young Curve-billedThrasher attendedby Brown Towhee. Auk 85:127-128. SVENSSON,S. 1955ß N6tviicka(Sitta europaea) sommatur Starungar (Sturnus vulgaris). Vfr Ffgelvfrld 14:256ß(abstract in Bird-Banding27:140ß) T,^XTER, B.A. 1930ß Father towheeadopts a familyßYosemite Nature Notes 9:88ß TICK•LL,W. L. N., ANDR. PINDER.1972. Chickrecognition by albatrossesßIbis 114:543- 548ß TRIVERS,R.L. 1971ß The evolutionof reciprocalaltruismß Q. Rev. Biol. 46:35-57ß TWOMBLY, F.D. 1934ß "Believe it or not." Bird-Lore 36:303ß VAN VELZEN,W.T. 1960ß Starlingsfed by Purple MartinsßAuk 77:477ß WARREN,H. M. 1930. The foster motheringof youngbirds. Bird-Lore 32:198ß W^RRINER,B.R. 1937ß Cardinalfeeds a youngcatbirdß Migrant 8:62ß WEINZIERL, H. 1958. Zwei weitere Falle von Mischbruten der Kohlund Blaumeise. Or- nithol. Mittß 10:31.(abstract in Bird-Banding29:196)ß WESTWOOD,R. W. 1946ß Contentsnoted. Nature Mag. 39:399ß WETHERBEE,MRS. K.B. 1930. Cooperativeparentsß Bird-Lore 32:202ß WItliT, E. M. 1934ß Attractingbirds at ChattanoogaßMigrant 5:46ß WILLIAMS,L. 1942ß Interrelationsin a nestinggroup of four speciesof birdsßWilson Bullß 54:238-249. WRItliT, H. 1956ß Towhee feeding behaviorßAID. Birdlife 4:27ß BiologyDepartment, Western Michigan University, Kalamazoo, Michigan 49008. Presentaddress: 2833 SpringDr., Fremont,Michigan 49412. Received20 Jul. 1981; accepted26 Mar. 1982. Vol53, No. 4 InterspecificFeeding [39 1

APPENDIXI. Common• and scientificnames of avianspecies mentioned in paperand list arrangedin taxonomicsequence (Morony et al. 1975).

Common name Scientific name

Arctic Loon Gavia arctica SpectacledEider Somateriafischeri Goshawk Accipitergentilis Red-tailed Hawk Buteojamaicensis American Kestrel Falcosparverius Herring Gull Larus argentatus Lesser Black-backed Gull Larusfuscus Common Sterna hirundo Ringed Turtle Dove Streptopelia risoria Mourning Dove Zenaida macroura White-winged Dove Zenaida asiatica Screech Owl Otus asio Great Horned Owl Bubovirginianus Green Violet-ear Colibri thalassinus White-earedHummingbird Hylocharisleucotis Red-bellied Woodpecker Melanerpescarolinus Williamson's Sapsucker Sphyrapicusthyroideus Downy Woodpecker Picoidespubescens Hairy Woodpecker Picoides villosus Northern Three-toed Woodpecker P icoidestridactylus Black-backedThree-toed Woodpecker Picoides arcticus Common Flicker Colaptesauratus Eastern Phoebe Sayornisphoebe Scissor-tailedFlycatcher Muscivoraf o•ficata EasternKingbird Tyrannustyrannus Great CrestedFlycatcher Myiarchuscrinitus Contopus virens LeastFlycatcher Empidonaxminimus Tree Swallow Tachycinetabicolor Purple Martin Prognesubis Cliff Swallow P etrochelidonpyrrhonota Gray Wagtail M otacilla cinerea Pied Wagtail M otacilla alba Bewick's Wren Thiomanesbewickii Carolina Wren Thryothorusludovicianus Winter Wren, Wren Troglodytestroglodytes House Wren Troglodytesaedon Gray Catbird Dumetella carolinensis Mockingbird Mimuspolyglottos Brown Toxostomarufum Curve-billed Thrasher Toxostoma curvirostre Hedge Sparrow Prunella madularis Robin Erithacus rubecula Redstart P hoenicurusphoenicurus Eastern Bluebird Sialia sialis Mountain Bluebird S ialia currucoides Townsend's Solitaire Myadestestownsendi Swainson's Thrush Catharus ustulatus Blackbird Turdus merula SongThrush Turdusphilomelos American Robin Turdusmigratorius Tropical P olioptilaplumbea 392] M.m. Shy j. FieldOrnithol. Autumn 1982

AVVE•D•X I. Continued.

Common name Scientific name

Willow Warbler Phylloscopustrochilus Karoo Prinia Prinia maculosa Layard'sTit-babbler P arisomalayardi Pied Flycatcher Ficedulahypoleuca SpottedFlycatcher Muscicapastriata Long-tailed Tit Aegithaloscaudatus Marsh Tit P aruspalustris Willow Tit Parus montanus Carolina Chickadee P arus car olinensis Mountain Chickadee Parusgambeli Coal Tit Parus ater Great Tit P arus major Blue Tit P arus caeruleus Plain Titmouse Parus inornatus Tufted Titmouse Parus bicolor Nuthatch Sitta europaea Pygmy Nuthatch Sittapyffmaea Certhiaf amiliaris Song Sparrow Zonotrichia melodia White-throatedSparrow Zonotrichia albicollis Dark-eyedJunco Juncohyemalis Dusky SeasideSparrow Ammodramus maritimus Chipping Sparrow Spizellapasserina Field Sparrow Spizellapusilla Rufous-sided Towhee Pipilo erythrophthalmus Brown Towhee Pipitofuscus Black-headed P heucticusmelanocephalus Cardinal Cardinalis cardinalis Scarlet Tanager Piranga olivacea Scarlet-rumpedBlack Tanager Rhamphoceluspasserinii Golden-maskedTanager Tangara nigrocincta Blue Honeycreeper Cyanerpescyaneus Black-and-white Warbler Mniotilta varia Blue-wingedWarbler Vermivorapinus Yellow Warbler Dendroicapetechia Pine Warbler Dendroicapinus Blackburnian Warbler Dendroicafusca Yellow-rumpedWarbler Dendroica coronata Kirtland's Warbler Dendroica kirtlandii American Redstart Setophagaruticilla Ovenbird Seiurus aur ocapillus Worm-eating Warbler Helmitheros vermivorus ProthonotaryWarbler Protonotaria citrea Kentucky Warbler Geothlypisformosa Yellow-breasted Chat Icteria virens Red-eyedVireo olivaceus Yellow-greenVireo Vireoolivaceus flavoviridis 2 Northern Oriole Icterusgalbula Red-winged Blackbird Agelaiusphoeniceus Common Grackle Quiscalusquiscula Chaffinch Fringilla coelebs Serin Serinus canaria Linnet Acanthis cannabina Vol. 53, No. 4 InterspecificFeeding [393

APPENDIX I. Continued.

Common name Scientific name

PurpleFinch Carpodacuspurpureus House Finch Carpodacusmexicanus Hawfinch Coccothraustescoccothraustes Zebra Finch Poephilaguttata BengaleseFinch Lonchurastriata House Sparrow Passerdomesticus Starling Sturnus vulgaris

• Common namesfor North Americanrecords follow the AOU Checklist(AOU 1957 and supplements). 2 Accordingto Peters(1931).

APPENDIXII. List of journals reviewed.Following the title of eachjournal listed is the dateof earliestvolume reviewed followed by volumeor issuenumbers. Parentheses indicate that the volumesenclosed were incompleteor missing.A dashbetween numbers indicates a continuousrun. A dash not followedby a number indicatesall volumeswere reviewed after the lastone listed. The literaturereviews were conducted in WaldoLibrary at Western MichiganUniversity, Kalamazoo, and theJosselyn Van Tyne Memorial Library at the Uni- versityof Michigan,Ann Arbor and reflecttheir journal holdings.

Alabama Birdlife. (1953)--l-- Iowa Birdlife. (1931)--1- Audubon(formerly Bird Lore). (1899)-- The Jack-PineWarbler. (1933)--13- 1--(12, 13)- The Kingbird. (1950)--1- The Auk. (1884)--1--(54)- The Loon (formerly The Flicker). (1929)-- Bird-Banding.(1939)--8• 1- Bird Lore. See Audubon. Maryland Birdlife. (1945)--(1, 4, 5)-- Bird Study. (1954)--1- The Murrelet. (1920)--1- The Blue Jay. (1942)--1- The Oriole. (1936)--1- The Cardinal. (1971)•l-- The Ostrich. (1968)--39-- The Chat. (1937)--1- The Raven. (1930)--1- The Condor. (1912)--11--(9, 10, 57- The Redstart. (1933)--1- 60)-- The Scissortail.(1951)--(1, 5-13)• The . (1968)--68-- The Wilson Bulletin. (1894)•1--(13, 20- The Flicker. See the Loon. 23)-- The Ibis. (1859)--1--