SELECTED DWIOHT VOCALIZATIONS OF THE COMMON CROW R. CHAMBERLAIN AND GEOROE W. CORNWELL BIO-ACOUSTICS has developedrapidly in the past decadeas a tool in the study of communicativemechanismsin wild birds (Hinde, 1969). The Common Crow (Corvus brachyrhynchos)is an ideal experimental subjectfor this type of research.Johnston(1961: 15) remarkshow even the casualobserverof crow behaviorcan detect changesin pitch and rate of call delivery, an indicationof the communicativepotential and behavioralcomplexityof crow vocalizations.In a study of the functional anatomyof the syrinx, Chamberlainet al. (1968) found the Common Crow to be morphologically capableof producinga significantvariety of notes. Bent (1946: 247-249) describesthe characteristic vocalizations of the Eastern CommonCrow (C. b. brachyrhynchos),citing observationsby Hoffman (1904), Knight (1908), Wright (1912), Allen (1919), Townsend (1923), and Forbush (1927). Allen (1919) relates crow notes to time rhythms, triplets, and other periodic combinations.Good (1952) discusses the meaningsof a numbero.f crow vocalizationsincluding: the rally call, given after sightingor hearinga crow predator; the squalling cry, uttered by a crow capturedby a predator; and the scoldingcall, given by adult crows during the nestingseasonwhen the young are threatened.The alarm, assembly,ordinary cawings,and beggingcries of the younghavebeentestedwith Americanand Frenchcrows,bothin Franceand the easternUnited States(Fringset al., 1958). Theseauthors tabulatedpositiveand negativephonotaxicresponses to. calls; they also tested crowswith the calls of other species,in many caseswithout observable effects on the crows' movements. Davis (1958) made audiospectrographic analyses. of typical calls of variousspeciesof North Americanand Mexicancrows,on the basisof whichhe describedthe crowof Sinaloa,Mexico,as a new species,Corvus sin.aloae.Chamberlain(1967) intensivelystudiedthe soundsproduced by the CommonCrowand the environmental and behavioralcontextin which each call occursin the wild. An annotatedmagnetictape of the vocalizatio.ns discussed in this paper is availablethroughinterlibrary loan of the thesisand from the Laboratoryof Ornithologyat CornellUniversity. Thompson(1969) showsthe cawingof CommonCrowsdisplaystwo kinds of organization,structuredand unstructured.He has evidencethat the rigid, idiosyncratic structuredpattern identifiesthe individualcrow; the unstructured patternreflectsthe degreeof the individual'sexcitement. On this basis,our paperdealsprimarilywith unstructured vocalizations, 613 The Auk, 88: 613-634. July 1971 614 CIClA3/iBERLAIN AIqDCORNWELL [Auk, Vol. 88 and only includessix sounds(contactcall, announcement call, duet notes, mimicry,courtshipsounds,and immaturehungerand feedingcalls) that may determineindividual recognition.We did not analyze crow vo.calizations for the stable and predictablecomponentsnecessaryfor species recognitionand distinctiveness. METItODS We recorded selected vocalizations AND PROCEDURES of wild and tame crows and noted the en- vironmental context and behavior accompa.nyingeach. Decoy mounted crows and Great Horned Owls (Bubo vlrginlanus) were used when appropriate to elicit calls requiring their presence. Selected crow calls and the distresscalls of three sympatric avian specieswere transcribed onto continuous loop tapes from the original field tapes. Loop tape tests were made from an automobile using a Wollensak 1700 T recorder with a 10-watt amplifier and a U.S. Army surplus cone speaker powered by a 12-volt car battery. The speaker was mounted on a ski rack on top of the automobile during field tests. Test tapes were broadcasted to crows in the field, with the automobile and testing equipment hidden as well as conditions allowed from direct sight of the crows. Crow responses to testswere classified,in part, by the following phonotaxicsystem derived from that of Frings et al. (1958): Class I: Specific predicted responseof crows within the estimated hearing range. Class II: Class III: Class IV: ClassV: Only part (quantified as much as possibleby field observation) of the crows within the estimated hearing range responded as predicted,while the remainderdid not respondat all. Only part of the crows within the estimated hearing range responded as predicted, while the remainder reacted in other than the predicted manner. No responseamong crows within the estimated hearing range. Unpredicted responsesby crows within the estimated hearing range. This ranking scheme was used for field classification of behavioral responsesto broadcastsof five crow vocalizations (Table 1). The expected responsewas predicted prior to broadcast. Actual responseswere compared to the predictions as a further confirmation of the context in which the vocalizations were recorded and tested. Vocalizations failing to elicit a responsewere tested repeatedly to establish the specific context under which they were given. The reactions to field tested vocalizationswere also classifiedas "positive" (q-) or "negative" (-) phonotaxis and "other" responses. "Positive" phonotaxis was recordedwhen crows respondedby flying directly to the broadcastsource. "Negative" phonotaxis was recorded when the responsewas a direct flight away from the broadcast source. The balance of responseswere tabulated under "other" (see tabular documentation in Chamberlain, 1967). Sounds that caused overt responsesamong crows were classified"primary" and nonresponsesoundswere designatedas "other." Three tape recorderswere used in the field to record Common Crow vocalizations: A Wollensak model 1700 T with a built-in converter and a Wollensak hi-impedance, omnidirectional, crystal microphone (50-8,000 c.p.s.) along with a Turner model 58, omnidirectional,hi-impedancemicrophone; a battery operated Kudelski Nagra III July 1971] Common Crow Vocalizations TABLE 615 1 CLASSIFICATION OF CROW I•.ES?ONSES TO CROW VOCALIZATIO]gS TESTEDAND I•.ANKED 1 Class of response Crow vocalization Prediction I Assemblycall Aggregationto sound source Simplescoldingcall Aggregationto 12 sound source 80% Dispersalcall Dispersalfrom 27 91% 8 sound source 67% Squallingcall Aggregationto sound source 22 88% Moribund call Aggregation to 5 soundsource 25% II 2 6% III 1 3% 3 20% 2 - IV V Total tests - - 30 - - 15 - - 1 - 3'0 8% - 25% 3 12% - - 25 2 7 20 4 10% 20% 12 10% 35% Extensive tabular documentationof crow responsesin Chamberlain (1967). One test included 21 Fish Crows. with a Nagra III omnidirectional microphone; and a Hitachi transistorizedportable model TRQ-370 with a Hitachi model TRQ-370 unidirectional microphone was used for short range recording. A 39-inch parabolic reflector was used to pick up distant crow vocalizationsand a 26-inch snow coaster disc to tape those nearby. Missilgrams made on a Missilyzer manufactured by the Kay Electric Company were used to illustrate call differences. PRIlVrARY VOCALIZATIONS As SElVrBLY CALL The assemblycall is an intenseand raucousmobbingcall containing long notes (wide band width of individual call notes) (Figure 1A), and is given when.crows see or hear nearby crow predators, e.g., the Great Horned Owl, Red-tailed Hawk (Buteo jamaicensis),or red fox (Vulpes ]ulva). The assemblycall has the apparent function of calling together a group.of crowsto drive predatorsaway from placesthe crowsfrequent, suchas nesting,roosting,or feedingareas. The CommonCrow assembly call attracts to. the sound sourceCommonand Fish Crows (Corvus ossi]ragus) within hearingrange (Table 1). Assemblycalls are given throughout the year. Field tests and/or observationswere made during all months o.f the year. Six night visits to Virginia crow roostsrevealedthat CommonCrows give assemblycalls at night. These night observationsare the basisfor all followingreferences to night vocalizations. When giving the assemblycall from a perchedand erectposition,the Common Crow holds the contour and wing feathers close to the body. The tail is retainedapproximately1 inch below the body plane, and the 616 CI-•A1VXBEIRLAIl• AI•TD CORIffVq'ELL [Auk,Vol.88 Figure1. A, assembly call; B, simplescolding call; C, modified scolding call; D, alertcall.(Certain vocalizations areillustrated morethanonce to show variation.) July 1971] Common Crow Vocalizations 617 rectricesspreadlaterally about 2 inchesbeyondthe tail's closedposition. The tail flicks up and down below the body plane during the call, and the wings move rapidly up and over and back and down to a closed position.Crowsemittingassembly callswhiledivingat an aerialpredator have a dihedralwing angle,and often extendthe legswith clawsopenin the mannerof a raptor closelypursuingprey. Crowsrespondingpositively to the assemblycall fly with rapid flappingwings. SIMPLE SCOLDING CALL The primary function of the simplescoldingcall (Figure lB) is to announcethe presenceand location of a crow predator. Good (1952) describesthis call. One of the three mobbingcalls the CommonCrow uses, the simplescoldingcall is more sharplystaccatoand continuousand not so raucousas the longer-notedassemblycall, althoughraucousnotesare commonto both calls. The simple scoldingcall is given when a crow hears an unseenpredator, and also while approachinga distant predator it can see. At close range the scolding vocalization is abandoned for assemblycalls,whichare moreeffectivein driving away a crowpredator. Simple scoldingcalls are deliveredat various degreesof intensity dependingon the type of predator,its distancefrom the crow, its movements, and the seasonof the year. For example, Common Crows give intenseand rapid scoldingcallswhenthey hear hootsof the Great Horned Owl, but less rapid and intensecalls when they hear the clucksof the Sharp-shinnedHawk (Accipiter striatus). Apparently this variation in calling occursbecausethe Great Horned Owl is a crow predator in winter roostsand springnestinggrounds,whereasthe Sharp-shinned Hawk rarely preys upon crows. Given throughoutthe year, day or night, CommonCrow simplescolding callsattra.ctboth Commonand Fish Crowsto the soundsource(Table 1). Crows do not remain near the test sourceof the simple scoldingcall for as long a time as with the assemblycall. They dispersed beforethe field testsendedin 5 of 15 broadcasts.Somebirds continuedto.repeat simple scoldingcallsup to 10 minutesafter field testsstopped. When giving the simplescoldingcall perchedand erect, the Common Crow movesthe head up and down and forward and backward. When the headmovestowardthe tail andup, the wingsflick up and away from the body. The beak closeswhen the head movesforward and down and openswhen the headmovesback and up. Crows used alternate flapping and gliding flight when approaching simplescoldingcalls during 4 and rapid flapping flight in 11 of 15 field tests. 618 MODIFIED C•^•r^• ^• Co•wtnn [Auk, Vol. 88 SCOLDING CALL The modifiedscoldingcall is a mobbingcall and is functionallysimilar to the simplescoldingcall, whichit closelyresembles, but all its notesare infiected(Figure 1C). It is utteredthroughoutthe year, day or night, in responseto the vocalizationor distant sightingof crow predators. Posturesaccompanying this call are nearlyidenticalto thoseof the simple scoldingcall. In five testscrowsdid not approachso rapidly or remain near the soundsourceso long as in response to.the simplescoldingcall. ALERT CALL The conversational alert call expresses the crow'sawareness of danger perceivedby auditory or visual means. Alert signalsvary in intensity, frequency,duration and number of notes (Figures 1D, 2A), and also vary in delivery accordingto. the immediacyand type of danger. For example,alert notesgiven by a ground-feedingcrow surprisedby a predator are sharp, intense,and rapid. Those uttered by a crow that seesa hunter approachingat a distanceare lesssharp,lessintense,and spaced at greaterintervals. The distinctivefeature that separatesthe alert call from simplescoldingis the numberof notesper call. The simplescolding call hasan unlimitednumberof notesper series,whereasthe alert sequence rarely has more than sevennotes, and generally six or less. The alert call is often deliveredmore rapidly than the scoldingcall. Alert calls did not provokea dispersalresponseamongcrowstested. Some birds indicate the type of predator with their alert calls, and some have specificcalls to. distinguishaerial from ground predators. Collias and Joos (1953) note that the hawk call of the Domestic Fowl (Gallus domesticus)is an intense,harsh scream,but that for a dog or man is a cackling sound. Catbird (Dumetella carolinensis) alert calls changeabruptly from short notes to protracted catlike "meows" as an intruder approachesthe nest (Collias, 1960': 369). We found no such differencesin crow alert calls, and 12 tests suggestthat the number of notesper crow alert seriesdoesnot conveyshadesof meaningor degree of danger. The function of the alert call is to warn crowsof nearby or distant danger.Crowsrespondto thissignalby emittingalert, assembly,modified or simplescoldingcalls,flying up into treesif on the ground,and flying away from and back to an arborealperch. Also, perchedcrowsshow nervousmovementssuch as rapid wing and tail flicking and frequent defecation. The posturesassociatedwith the alert call are similar to thosefor the simplescoldingcall. In 12 of 17 tests,crowsresponded positivelyto the alert call by givingassembly and simplescoldingcalls whileapparentlysearching for the nonexistent predator. July 1971] DISPERSAL Common Crow Vocalizations 619 CALL The dispersalcall (Figures 2B, 2C) is the extremedegreeof the alert call. Its notesare generallysharper(thinner band width), and they are deliveredclosertogetherthan alert notes. The first note is often infiected (Figure 2B). The variability in soundintensity betweenalert and dispersalsignalswas not measured, althoughwe suspectthat dispersalcalls are deliveredwith greater volume than alert calls. The. dispersalcall made CommonCrows fly directly away from the test area during eight broadcasts.In three testsCommonCrowscameto the call giving mobbing callsand apparentlylookingfor a predator (Table 1). Dispersalcalls are given by crowsthat senseimmediatedanger,such as a hunter'ssuddenappearancenearby. Crowsin roostscommonlygive oneafter sightingdanger,say a Great HornedOwl. The numberof notes in a dispersalcall doesnot appearto conveydifferent degreesof danger to responding crows.We believethis is the vocalizationFringsand Frings (1957) refer to as the alarm call consisting of "four short notes." We foundthe dispersalcall to vary in notenumber. In eight testsof dispersal calls scoresof crows rose out of trees and flew high in the air, massing in a circlingflock and giving simplescoldingand assemblycalls. Frings and Frings (1964: 58) state, "If a crow suddenlyflies away from the group without making any sound,the. others usually follow. It has spreadan alarm--the fastestand safestway." Chamberlainmade a supposed"sentry" perchedabove a flock of ground-feedingcrows fly on four different occasionsduring this study without giving a dispersal call. In each case the flock remained on the ground until Chamberlain camewithin their sight. SQUALLING CALL Squallingca]Isare givenby crowsstrugglingto.escapefrom a predator suchas a hawk or fox, and can be producedby holdingan immatureor adult crowupsidedownby its legsand shakingit (Figures2D, 3A). It variesin intensity•frequency,tempo,and durationwith the gravity of the situation. For example,a crow that is shakenvigorouslydelivers squallingnotesmore intensive]y,sharp]y,and rapidly than one shaken slowly. Squallingca]Isare generallyhigherpitchedand have more of a wailingquality thanothercrowvoca]izations. Common and Fish Crows respond to immature and adult Common Crow squallingca]Isby gatheringrapidly to the soundsource(Table 1). At night CommonCrowswi]] fly to squallingca]Is near a roost. One Januarynight in westernVirginia, crowsgaveassemblyand simplescolding ca]Is over the soundsourcebefore settlingback into coniferssome 620 CI•A1VZBERLAIN A•D CORNWELL 0.5 LO TIME IN 05 IN L5 • SECONDS I0 TIME [Auk, VoL 88 15 20 SECONDS c 10 TIME iN SECONDS TIME IN SECONDS t5 20 1,5 20 i D I0 Figure 2. A, alert call; B, dispersalcall; C, dispersalcall; D, squa111ng call. July 1971] CommonCrow Vocallzatlons 621 A 015 LO TIME 0.5 I• go I•i gO IN • LO c Figure3. A, squallingcall; B, squallingcall broadcasted in reverseat the speed originallyrecorded; C, moribundcall; D, threat call. 622 CHA1V•BERLAIN ALOUD CORNWELL [Auk, Vol. 88 200 yards from thoseoccupiedbeforethe test. Broadcasting distresscalls of the EuropeanJackdaw(Corvusmonedula)eliciteda negativeresponse at night in Frenchcrowroosts,causingbirds of severalspeciesto desert the roostspermanently(Fringset al., 1958). Crows respondto squallingnotes with assembly,modified, and simple scoldingcalls. They apparentlyapproachedthe soundsourceintending to mob the crow predator. We broadcasted adult squalling calls in a crow roost in westernVirginia in January with the tape reversed. The crows respondedby aggregatingto the sound sourcein spite of tape reversal (Figure 3B). MORIBUND CALL The moribund call (Figure 3C) is the extreme squallingcall of the CommonCrow and is emitted by a dying crow. Moribund calls are not deliveredas rapidly or loudly, but are more raucousand gurglingthan squallingnotes. Becauseof its weakenedconditionwhile giving this call, a crow doesnot struggleor fight off a predator. The moribundcall is a definite communicationproducedby the dying bird and not a sound producedmechanicallyor incidentallyin the processof dying. Crows showed variable responsesto the moribund call tested in the field. In 20 tests, crows exhibitedpositive phonotaxisin 5, negative phonotaxis in 7, and made8 other responses (Table 1). THREAT CALL The threat call (Figure 3D) is the mostvariableof the CommonCrow's vocalizations.Threat calls are often given as screams,cackles,staccato and rattling notes,coos,and a variety of other sounds. They occur in a highly variablecontextand are often indescribable.A crow may utter one when under attack or threat, or when attacking or threateninga predator or another crow. Crows respondedto five field tests in an unpredictable manner. Threat calls are the commonestnotes heard among roosting Common Crows. The birds compete with one another for roosting perches. In- comingcrowstry to.dislodgethosealreadyroosting,and both the attackers and those attacked utter threat calls. The apparent function of these signalsis to make the opponentretreat. Good (1952) describesgrowlcalls givenby CommonCrowsdiving or pursuinga wingedpredatorat closerange. Prior to diving at a flying predator,a crow will first hover on beatingwings,then start an erratic descentwith wingsclosedin a dihedralover the rump. In sevenobservations the legs were extended with open claws, and a type of threat July 1971] Common Crow Vocalizations 623 ID Figure 4. A, growl call; B, immature hunger and feeding call; C, contact call; D, announcement call. 624 CItAMBERLAIiN ^•) COR•WE• [Auk, Vol. 88 vocalization (or growl call) was emitted when the crow passedclose to the predator (Figure 4A). Crows while mobbing a flying or perched raptor give assembly,simpleand modifiedscoldingcalls, but supplant them with threat calls when close to or within striking distance of a raptor. One April Chamberlainheard an adult CommonCrow emit threat signalswhen diving upona Turkey Vulture (Carthartesaura) that came near its nest containingthree fledglingsin west central Florida. During threat vocalizationwhile perched,the neck, nape, throat, and head feathersare erect and appear ruffled. The wings move back and up when the bird rears back while under attack of when threatened. The wingsmay be droopedand spread,and the headloweredbetweenthe legsprior to attackingor beingattacked. IMMATURE HUNGER AND FEEDING CALLS Immature CommonCrowsgapeand give thesecalls (Figure 4B) when stimulatedby hunger or the sight of food, by seeingor hearing objects associated with food,or by a combinationof any or all of these. Hunger calls elicit variable responsesamong crows in the field. Frings et al. (1958) statesthat "beggingcriesof youngnestlings"had no observable effect on the movements of crows when field tested. In 31 tests we obtained 14 positive,9 negative,and 13 other responses to hunger calls broadcastedin the field (8 tests showedtwo classesof responses). The hungercall resembles. squallingnotes,but its individual notesare no.tso protractedand wailing. The feedingcall containsa numberof rapid but clearly distinguishable notesdeliveredby the youngwhile swallowing food after uttering hunger notes while gaping. Hunger calls are subject to differentdegreesof delivery,dependenton the amountand proximity of food in relation to the calling bird. The stronger the stimulus, the fasterand moreintensethe delivery of the high-pitchednotes. Muir (1954) noted that unfed young Tawny Owls (Strix aluco) call more often than fed ones,thereby expressingtheir degreeof hunger; he suggests this behaviormay bring about a more even food apportionment amongthe youngby adult owls. We notedthis samephenomenon in both tame and wild CommonCrows. Immature hunger calls also function as locationnotes,helpingparent crowsreestablishcontactwith their young. Pet crows often associate food with a human voice when talking ac- companieshand feeding; these.birds stop their insistent hunger calls when the ownersare concealedand quiet, but promptly resumegaping and the accompanying vocalizations whenthey seeor hear their owner. Many bird speciesare known to have food calls given by the adult that stimulatefeedingresponses in the young,suchas the YellowWarbler July 1971] Common Crow Vocalizations 625 (Dendroica petechia) and Least Flycatcher (Empidonax minimus) (Collias, 1960: 369). We detectedno specificvocalizationgivenby parent crowsto their young that could be interpretedas a food call. Nestling crows respondwith hunger calls to virtually all vocalizationsof adults near the nest,includingthe assembly,simplescolding,alert, and dispersal calls. While giving hungercalls in or out of the nest when perched,immature CommonCrows move their wings rapidly up and down away from the body. The tail alsomovesup and down during begging. The bright-red mouth is evidentduring the period of open-mouthedfood begging. CONTACT CALL The componentsof group movementsby CommonCrows are assembly, preparationfor movement,initiation, liaison while moving, and cessation. Land birds that travel in flocksappearto placeheavy relianceon contact notes betweenindividuals. Marshall (1964) lists a "seep note" by the Brown Towhee (Pipilo fuscus)that mated birds use to maintaincontact with each other when they are a short distance apart but hidden from eachother. Odum (1942) notesthat the "chick-a-dee-dee-dee" call of the Black-cappedChickadee(Parus atricapillus) is usedto maintain contact when the flock is scattered. Contact notesthat function to keep the flocks togetherare commonin migrating crows (Figure 4C). This call char- acteristicallycontainsthree, four, or more notes;but, commonlya fournoted sequenceis uttered by one crow in note pairs. CommonCrows maintaingroupcohesion by answeringeachotherwith identicalor similar contactnotes,particularlyin roostsand in migratingflockswhen pairs become separated. Crows emit contact calls while in flight or perched. When perching, the associatedposturesare similar to thoseusedin simple scoldingcalls. ANNOUNCEMENT CALL Announcementcalls (Figure 4D) are used by individualsflying to join an aggregationof CommonCrows, such as those at feeding grounds or roostsand functionas a form of recognitionand contact. These calls are extremelyvariable in quality of delivery, and often closelyresemble simplescoldingand assemblycalls,but are usuallymuchshorter. Common Crowsgive thesecalls when in gliding or flapping flight. During gliding flight the bird descendsat approximately45 degreeswith wings held in a dihedral position,the tail level with the body or angled dorsally. On five instances, an answerwasreceivedfrom a flockmember(s) beforethe bird resumednoisy,flappingflight. 626 DUET C•^:•BERL^• * AI•IDCORNWELL [Auk, Vol. 88 NOTES These notes (Figure 5A) were heard betweentwo tame, caged adult CommonCrowsseparatedby an opaquepartition. Both birds appeared to be males. These two crowsdevelopedfour recognizableduet combinations including rattling notes and cawing variations that they used to establishmutual recognitionwhen out of each other's sight. Apparently paired wild crowsalso use these duets for individual recognitionand contactin roostsor largefeedingflocks. Many bird speciespractice duetting,generallybetweenpairs (Thorpe and North, 1965). CommonCrows are no exception: one crow emits a certain vocalization that stimulates another crow to answer in kind. Five duetswere heard betweenfive different pairs of wild crowsin roosts during this study. COURTSHIP VOCALIZATIONS The observation and recording of vocalizations associatedwith pre- copulatorybehavior in the Common Crow were too limited to justify reportinghere. Bent (1946: 227-229) describes the behaviorand vocalizationsaccompanyingCommonCrow mating. OTHER VOCALIZATIONS Juvenile notes.--Immature crows in their first year have a variety of juvenile notesthat are apparentlypracticedwhile gainingexperienceprior to actual use (Figure 5B). An immature Common Crow Chamberlain heard in July emitted juvenile.notes resemblingassemblycalls. In west central Florida we recordedan immature Common Crow in April trying to give the screamspeculiar to the Florida population of the Common Crow (C. b. pascuus) (see below). Juvenile notes are given with the head loweredand beak open. The wings flick up and away from the body and down to a closedposition. Contentment no.tes.--Five tame crows in southwesternVirginia gave contentmentnoteswhen they had fed to capacity,were having their throat feathers stroked, or were otherwiseapparently content. These notes are soft and plaintive to the ear (Figure 5C). While emitting them the bird sits low on a perch in contrast to the erect position during the alert call. Rattling notes.--Rattling notes are heard within crow roosts as well as from flocksof feedingand migratingcrows(Figure 5D). This vocalization generallycontains5-20 sharply delivered,staccatonotesin a single utterance, somewhat like a rapid woodpeckertapping. Rattling notes are given by apparently paired crowswhen diving near each other, while descending into a roost,or whenenteringa diurnal feedingflock. Clucking July 1971] Common Crow Vocalizations 627 A B c 0.5 I:O L5 ?•0! Figure 5, A, duet notes; B, juvenile notes; C, contentmentnotes; D, rattling notes. 628 Ca^•BERL^•S ^•m CORNWEL• [Auk, Vol. 88 sounds,intermediatebetweenrattling and threat notes,are utteredwithin feedingand migratingflocks. In July in southwestern Virginia, a juvenile crowgavehungercallswhile approaching an adult crowin a tree, to which the adult respondedwith rattling notes..Given by both male and female CommonCrowsin captivity, rattling noteselicitedno observableresponses from wild crows when field tested. Rattling notesmay be oneof the male courtshipcallsaddressed to.the femaleprior to matingin the spring. The femaleanswerswith submissive vocalizations similar to thoseimmatureCommonCrowsgive when begging for food (Townsend,1927). Townsendstatesthat rattling notesmay also be emittedby malescominginto sexualmaturity for the first time in their secondfall while aerially pursuing females. One Septemberin western Virginia, Chamberlainwatchedat least 14 CommonCrows of unknown sex and age erratically pursuingand diving upon each other while constantly givingrattling and threat calls. Wow-wow notes.--•Vow-wownotes (Figure 6A) were given repeatedly by three adult tame crowsbut were never heard from crowsin the wild. This sequencesoundslike a dog yelping, but was not believed to be mimicry becauseit was heard from birds in New York and Virginia. No function could be associated with this vocalization. While giving wow-wownotes,the head is moved up and down as if bowing,thebeakis opened andclosed, thetail feathersarespreadlaterally, the wingsmoveup and away from the body and downto a closedposition, the nictitatingmembranecoversthe cornea,and the bellyshakesnoticeably. Carr-carr notes.--Carr-carr notes are protracted caws (Figure 6B). These were recordedin Septemberat a crow roost in western Virginia. We saw no responseto them by roostingcrows. We twice heard threesyllabledcarr-carr notes uttered near an active crow nest by an adult crowin May, and onceby an adult crowin a flock of eight crowsperched in elm trees in September,both in western Virginia. No responseor particularcontextwasnotedin eitherinstance. Whisper notes.--•Vhisper notes, as implied, are muted. We recorded an adult crow giving whisper notes (Figure 6C) resemblingalert calls in March in western Virginia. Staccato whisper notes also were heard in a crow roost one February in southwesternVirginia. No responsewas noted in either instance. Coo notes.--In Myakka River State Park, Florida, Chamberlainwalked near a family group of CommonCrowsin April. The two adults uttered assemblyand simplescoldingcalls,and one adult gave a two-notedhigh pitched "coo-coo" (Figure 6D) between these mobbing calls. Nearby crows made no. apparent response. A coo note was again heard in the July 1971] Com•non Crow Vocalizations 629 A B D Figure 6. A, wow-wow notes; B, carr-carr notes; C, whisper notes; D, coo notes. 630 C•^•B•L^X• A•D CO•t•WEL• [Auk, Vol. 88 s' A 0,5 0,5 Figure7. A, organnotes(delineated by arrowsnext to a scoldingnote); B• wah-oo notes; C. C. b. pasctatsscreams. samelocale,givenby an adultcrowpriorto an alertcall directedat the observer.Coonotesmay be associated with danger.Similarsounds have beenheardfrom tame crowswhenseeingstrangepeople,dogs,etc. Townsend(1927) describes apparentlysimilarcoonotesas one of the male precopulatoryvocalizations. Organnotes.--Chamberlain useda crowcallto lurecrowsto a mounted Great Horned Owl in westernVirginia in March. The crowsgavesimple scolding andassembly callswhilemobbing thedecoy.Duringthemobbing• July 1971] Common Crow Vocalizations 631 one crow with a flock perchedin a clusterof conifersuttered at least 10 high-pitched organnotes(Figure 7A). No particularresponse by individual crowsperchednear the crow uttering organ noteswas seen. Organ notes may be similarto coonotesin that they are associated with danger. Wah-oo note.--In southwesternVirginia a crow was recordedgiving a singlewah-oo note in a roost in February approximately10 minutes before sundown (Figure 7B). On 12 occasionsduring the study, crepuscularcrow sounds similar to wah-oo notes were heard, but not taped. Roostingcrows emit many soundsnot encounteredelsewhere, includingsoundsassociatedwith threat. For example,one adult crow in a flock of crowsflying into a roostin westernNew York uttered highpitched,four-syllabledchippingnotes. C. b. pascuusscreams.--Individualsin the Florida population of the CommonCrow commonlyproduceintense,raucousscreams(Figure 7C). Crows near the sound source make no apparent response. Bent (1946: 262) statesthat severalobservers havereporteda vocalization peculiar to the Florida Crow. Mortimer (1890) heard soundslike the criesof a cuckoo(Coccyzussp.). These may have been similar to the crow screamswe heard in Myakka River State Park, Florida. Mimicry.--Common Crows often mimic other birds and mammals. Forbush(1927) hearda crowgive an excellentimitationof the whine of a dog. He also heard crows emit a varied assortment of notes, some of which were imitations,suchas the cry of a child, the squawk of a hen, and the crow of a youngrooster. In captivity, crowslearn to mimic the human voice, as Chamberlain noted in eight tame crows studied. One adult male crow in western Virginia, couldsay "oh my God, oh Lord" very clearly. An adult male crowin westernNew York couldsay "hello" loudly and clearly. One in Florida announced"I'm Jim Crow." The myth regarding splitting a crow'stonguebeforeit will "talk" is probablyattributable to the natural split at the tip of the tongue. One of the functionsof mimicry may be to establishand strengthen the individualpair bond (Thorpe and North, 1965). This is particularly so in those specieswhere territorial aggressiveness is not very marked, as the Common Crow (Good, 1952). In certain species,this ability enables each bird to learn the vocal contribution of its mate as well as its own (Thorpe and North, 1965). Crowsmay usethis trait in maintaining contactwith a mate. If so, the extremeof this unusualimitative ability of birdsin parrots(Psittacidae),Starlings(Sturnusvulgaris),and crowsis plausiblyexplained. PerhapsCommonCrows in the wild use thesepowersto imitate the idiosyncrasiesand inflectionsin the vocalizationsof their mates,or othermembersof a flock. 632 Ct:IA3.•BERLAIN AND CORNWELL [Auk, Vol. 88 Interspecificresponseto distresscalls.--CommonCrowswere testedby distresscalls of the Starling, Bluejay (Cyanocitta cristata), and the CommonGrackle (Quiscalusquiscula). They gatheredto the Bluejay calls in 5 of 10 tests,showedno responseto the CommonGracklecalls in 5 of 6 tests,and were unpredictablein responseto the Starlingcalls. Fish Crowsrespondedpositivelyin all testsof the Starling call. ACKNOWLEDG2V[ENTS Release 68-1 of the Virginia Cooperative •Vildlife Research Unit, Virginia Commission of Game and Inland Fisheries,Virgina Polytechnic Institute, Wildlife Management Institute, and U.S. Fish and Wildlife Service,cooperating.This investigationwas financed by the Frank M. Chapman Memorial Fund and The Virginia Commission of Game and Inland Fisheries. The authors thank S. A. Cable for technical assistance and equipment maintenance and for permitting use of the electronic equipment in the V.P.I. Radio and Motion Picture Studio. Appreciation goes to Lee Wilkins of Deerfield, Virginia, for encouragement, maintenance of electronic equipment, and helpful technical advice. We especially thank Peter Paul Kellogg, Miss L. V. Engelhard, and the staff of the Library of Natural Sounds, Cornell University for the use of the Missilyzer and other electronicequipment. We are most grateful to Mrs. Sadie Wolvin and to the others who let us record the vocalizations of their tame crows. We thank the staff of the Myakka River State Park, Florida for allowing us to record crows on the premises. We are indebted to Mrs. Anne Tatlow IV, Gary Sale, Robert Oster, Gary Howard, Glenn R. Dudderar and the V.P.I. Graduate students who aided in the field testing of crow calls during this study. SUMMARY This study documents23 of the vocalizationsof the CommonCrow, and describesthe appropriateenvironmentaland behavioralcontext for each. Ten calls were tested in the field to further substantiate and define predictedcrow responses to thesesignals. Crows respondedto nine of these field-testedcalls. Crow vocalizationsdescribedare: assemblycall, simple scoldingcall, modified scolding call, alert call, dispersal call, squallingcall, moribund call, threat call, immature hunger and feeding calls,contactcall, announcement call, duet notes,juvenilenotes,contentment notes,rattling notes,wow-wownotes,carr-carrnotes,whispernotes, coonotes,organnotes,wah-oonotes,C. b. pascuusscreams,and mimicry. Crowresponses to interspecific distresscallsare noted. Thesevocalizations are not all the sounds of the Common Crow known to the authors or other workers. The vocal repertoireof CommonCrowsshowsconsiderable diversificationand specializationin relation to behavior patterns concernedwith flockingandwith predators. An investigation to determine which sounds are innate, learned, or combinationsof both shouldfollow the completedocumentationof Common Crow vocalizations.Indeed, the study of sound variation in passefine July 1971] Common Crow Vocalizations 633 populationsis important for the analysisof developmental, genetic,and comparativeaspectsof vocalizationas a biologicalphenomenon. LITERATURE ALLEN,F. H. 36: CITED 1919. The aestheticsensein birds as illustrated by the crow. Auk, 112-113. BENT, A.C. 1946. Life historiesof North American crows,jays and titmice. U.S. Natl. Mus., Bull. 191: 227-229, 247-249,262. C>tAMBERLA•N, D.R. 1967. The vocalizationsand syringealanatomy of the Common Crow, Corvus brachyrhynchos. Published M.S. Thesis. Blacksburg, Virginia, Virginia Polytechnic Inst. (Available from University Microfilms, Inc., Ann Arbor, Michigan.) CIIA1VfBERLAIN, D. R., W. B. GROSS,G. W. CORNWELL,AND H. S. MOSBY. 1968. Syringealanatomy in the Common Crow. Auk, 85: 244-252. COELtAS, N. E. 1960. An ecologicaland functional classificationof animal sounds. Pp. 368-391 in Animal sounds and communications (W. E. LaDyon and W. N. Tavolga, Eds.). Washington,D.C., Amer. Inst. Biol. Sci. COLL•AS,N. E., AND M. Joos. 1953. The spectrographicanalysis of sound signals of the Domestic Fowl. Behavior, 5: 176-188. DAWS, L. I. 1958. Acoustic evidence of relationship in North American crows. Wilson Bull., 70: 151-167. FORBUSH, E.H. 1927. Birds of Massachusetts and other New England states. vol. 2. Norwood, MassachusettsDept. Agr. FRINGS,H., ANDM. FRINGS. 1957. Recorded calls of the eastern crow as attractants and repellents. J. Wildl. Mgmt., 21: 91. FRZNGS,H., AND M. FraNCS. 1964. Animal communication. New York, Blaisdel Publ. Co. FmN6s, H., M. FRINGS,J. JUMBER,R.-G. BUSNEL,J. GIBAN, ANDP. GRAMET. 1958. Reactions of American and French species of Corvus and Larus to recorded communication signalstested reciprocally. Ecology, 39: 126-131. GOOD,E.E. 1952. The life history of the American crow--Corvus brctchyrhynchos Brehm. Unpublished Ph.D. Dissertation, Columbus, Ohio State Univ. H•qDE, R. A. 1969. Bird voca]izations. New York, Cambridge Univ. Press. HOFFMAN,R. 1904. A guide to the birds of New England and easternNew York. Cambridge,Massachusetts,Riverside Press. JOHNSTON,D. W. 1961. The biosystematics of American crows. Seattle, Univ. Washington Press. KNIGHT, O. •V. 1908. The birds of Maine. Bangor, Maine, C. H. Glass Co. MARSHALL, J. 1964. Voice in communication and relationships among Brown Towhees. Condor, 66: 345-356. MORTIMER,D. 1890. Notes on habits of a few birds of Orange County, Florida. Auk, 7: 337-343. MUIR, R.C. 1954. Calling and feeding rates of fledged Tawny Owls. Bird Study, 1: 111-117. ODUM, E.P. 1942. Annual cycle of the Black-cappedChickadee. Auk, 59: 499-535. T}rOMPSON,N.S. 1969. Caws and affect in the communicationof Common Crows. Bull. Ecol. Soc. Amer., 50: 142. T>tORPE,W. H., ANDM. E. W. NORT>t. 1965. The lonely tunesmithsof nature-men and the birds. Saturday Rev., 68: 85-87. 634 CttAMBERLAIN AND CORNWELL [Auk, Vol. 88 TOWNSEND,C.W. 1923. The voice and courtship of the crow. Bull. Essex County Ornithol. Club, 5: 4-8. TOWNSEND,C. W. 1927. Notes on the courtship of the Lesser Scaup, Everglade Kite, crow, and Boat-tailed and Great-tailed Grackles. Auk, 44: 549-554. WRIGttT, H. W. 1912. Morning awakening and even-song. Auk, 29: 307-327. Department o• Zoology, University o7 Maryland, CollegePark, Maryland 20742, and Schoolof Forestry, University of Florida, Gainesville, Florida 32601. Accepted30 May 1970.
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