Mobbing Behavior of a Pair of Elegant Trogons Author(s): Jack F. Cully, Jr. Source: The Condor, Vol. 88, No. 1 (Feb., 1986), pp. 103-104 Published by: University of California Press on behalf of the Cooper Ornithological Society Stable URL: http://www.jstor.org/stable/1367766 . Accessed: 27/05/2011 12:34 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=ucal. . Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. University of California Press and Cooper Ornithological Society are collaborating with JSTOR to digitize, preserve and extend access to The Condor. http://www.jstor.org 103 SHORT COMMUNICATIONS The Condor 88:103-104 ? The Cooper Ornithological Society 1986 MOBBING BEHAVIOROF A PAIR OF ELEGANTTROGONS1 JACK F. CULLY, JR. New Mexico 87131 USA Departmentof Biology, Universityof New Mexico, Albuquerque, Key words: Elegant Trogon;GreatHornedOwl;mobbing;move-onhypothesis;predatordeterrence. Mobbingbehavior among birds is considereda mechanism of predatordeterrence;however,the benefitsof mobbing to small or passively mobbing species is not clear (Cully and Ligon 1976, Curio 1978). Most species mob by giving loud calls combined with wing and tail flitting that, together,appearto attractadditionalbirds of many species. I report here my observationson the mobbing behaviorof a pairof ElegantTrogons(Trogonelegans)to a GreatHornedOwl (Bubovirginianus).Althoughthe trogons maintained a substantialdistance between themselvesandthe owl,theircallsquicklyattractedotherspecies, some of which mobbed very aggressively.By attracting more aggressivespeciesto mob, small birds may be able to have predatorsdriven away from their territoriesand derive the same benefits as the more aggressivespecies (the "move on" hypothesisof Curio [1978]). On 4 July 1970 at the South Fork Campgroundin the ChiricahuaMountains, ChiricahuaCounty, Arizona, I tethereda live GreatHornedOwl 60 cm above the ground at the base of a small tree. It was immediatelymobbedby a pair of ElegantTrogons.(I laterdiscoveredthat the tree containedtheir nest.) I observedthese birds for approximately 30 min beforeI terminatedthe sessionby moving the owl away from the area. After the owl was removed the trogonsfreely enteredtheir nest. I watchedthe birds from behind some shrubsat a distance of about 10 m. Calls were recordedon a Uher 4000 Report-Ltape recorderat 3.5 IPS using a Uher M514 microphone.The calls were analyzed on a Kay Electric Company Sonagraph6061B. I did not attemptto count calls or flits, and I did not attemptto measurethe durationof the response. Within 1 min after I exposed the tetheredowl, a male trogonbeganto mob from a distanceof about 15 m. He was perchedon a branchapproximately5 m high in full view of the owl andme. Within2 min afterthe maletrogon beganmobbinghe wasjoined by his mate. Althoughtheir calls wereloud and attractedvariousotherbirds,no other trogonsjoined the mob. The male trogon flew to a branch in view of the owl about 15 m away. He called and occasionallyflitted his wings and tail. There was very little jumping or flying betweenperches.Whenthe femalearrivedshetook a perch with more cover about 6 m from the owl, and also began to call.Bothtrogonschangedtheirperchesduringthe mobbing session, but they never approachedthe owl closer than 6 m. ElegantTrogonsuseda wing-tailflitwhilemobbing.The trogonsfirstrapidlyflickedthe wings (durationabout 0.1 sec), and then slowlyraisedand loweredthe tail (duration approximately1.0 sec). Therewas no bowingof the body associatedwith the flit. Trogonmobbingcalls consistedof a rapidseriesof loud staccatochucksgiven in phrasesof 4 to 10 notes (Fig. 1). The male'scalls soundedhigherpitchedthanthe female's, but otherwisethey weresimilar.Harmonicsrangedin frequencyfrom 1.0 to 2.5 kHz in the male and from 0.5 to 6.0 kHz in the female. The dominant harmonic of the male was centeredat 1.5 kHz, that of the female at 0.8 kHz. Since high-frequencyharmonicsattenuatefasterin woodlandsthanlow frequencies(Martinand Marler1977), the shorterfrequencyrange recordedfor the male may have resulted from my recordinghis calls at a greater distance.Mobbingcallphrasesrangedfromapproximately 0.5 to 1.2 sec in duration.The individualnotes were regularlyspacedat 0.1 to 0.12 sec intervals.Eachnote of the phrasewas of similar duration(0.08 sec) and frequency range.The energyof the calls was given in a rapidburst. The frequencyrange and amplitudecharacteristicsof the calls of many speciesof mobbingbirdsarewell suited to facilitatelocatingthe callingbirds,which makesit easy for more birds to join the mob (Marler1955, Marlerand Hamilton 1967, Martin and Marler 1977). The calls of ElegantTrogons are similar. High frequenciesattenuate quickly,makinga wide frequencyrangeuseful for determiningthe distanceto the source(MartinandMarler1977). The calls were repetitiveand beganand ended abruptlycharactersthat enhanceddirectionfinding. Trogonmobbingwas passivecomparedto that of many otherspecies.Gray-breastedJays(Aphelocomaultramarina), which joined the mob within 1 min after the male trogon began calling,jumped and flew constantly,often diving within 5 cm of the owl's head. Passerinemobbing is characterizedby loud calls, numerousjumps and flights from perch to perch,and frequenttail flits (Hinde 1954; Cully and Ligon 1976, in press).In passerines,the tail flit Male 4 0 4' A I i.. N I 6 A " Female 4- 0 I I I ' I .. 0 Time In Seconds ' Received 18 March 1985. Final acceptance7 September 1985. . ' .. .I . ............. i ' ..I. 1 FIGURE 1. Sonogramsof male and femaleElegantTrogon mobbingcalls. 104 SHORTCOMMUNICATIONS usedin mobbingprobablyaroseas a flightintentionmovement (Daanje 1950, Hinde 1954). A component of the evidence for its origin as a flight intention movement is the changein posturethat accompaniesthe wing and tail movements;the movementis similarto theposturalchange that often precedesflight.This componentof the flit (the change in posture)is missing from trogon mobbing tail flits. This suggeststhat in mobbingthe importantfactor is movement,whichenhancesvisibilityto otherbirdsand perhapsconfusionof the predator(but see Curio 1978). Trogon mobbing may illustratethe value of mobbing among small birds or other passively mobbing species. Althoughthese species are not likely to drive largepredators from their territories,their calls often attractother, more aggressivebirds such as jays. Oftenwhen I placeda captive owl on a perch to observethe responsesof other birds, the first species to mob were smallerspecies such as PlainTitmice(Parusinornatus)or Black-throatedSparrows (Amphispizabilineata). When jays arrived they mobbed aggressively;on several occasions they startled the tethered owl sufficientlyto cause her to fall off her perch.This suggeststhat mobbingmay benefitthe smaller speciesby atttractinglargerbirdswho can forceowls, particularysmall species, to flee. I thank J. D. Ligon, J. Wiens and B. Woodwardfor reviewingdraftsof this paper. LITERATURECITED CULLY, J. F., JR., AND J. D. LIGON. 1976. Comparative mobbingbehaviorof Scruband Mexicanjays. Auk 93:116-125. CULLY, J. F., JR., AND J. D. LIGON. In press. Seasonal factors in mobbing intensity of the Pinyon Jay. Z. Tierpsychol. of avianmobCURIO, E. 1978. The adaptivesignificance bing. I. Teleonomic hypothesesand predictions.Z. Tierpsychol.48:175-183. DAANJE, A. 1950. On locomotorymovements in birds andthe intentionmovementsderivedfromthem.Behaviour3:48-98. HINDE,R. A. 1954. Factors governingthe changesin strengthof a partiallyinbornresponse,as shown by the mobbing behaviour of the Chaffinch(Fringilla coelebs).I. The nature of the response and an examinationof its course.Proc.R. Soc. Lond.Biol. Sci. 142:306-331. of some animalcalls. MARLER, P. 1955. Characteristics Nature 176:6-8. III. 1967. MechaMARLER, P., ANDW. D. HAMILTON nisms of animalbehavior.Wiley, New York. 1977. Soundtransmission MARTIN, K., ANDP. MARLER. and its significancefor animal vocalization.Behav. Ecol. Sociobiol. 2:271-290. TheCondor88:104-105 ? The CooperOrnithologicalSociety 1986 SPITEFULNESS,ALTRUISM, AND THE COST OF AGGRESSION: EVIDENCE AGAINST SUPERTERRITORIALITY IN TREE SWALLOWS' RALEIGH J. ROBERTSON, H. LISLE GIBBS2 AND BRIDGET J. STUTCHBURY Departmentof Biology, Queen'sUniversity,Kingston,OntarioK7L 3N6, Canada For such spitefulnessto evolve, the benefitsto the suKey words: Superterritory;aggression; hole-nesting; TreeSwallow. individualmust exceedthe costs. The excluperterritorial sion of a potential breederby an inhibitoryindividual resultsin an equalincreasein relativefitnessamongstall is a spiteful behavior in which an in- breeders,whetheror not they inhibit others.In contrast, Superterritoriality dividual'srelativefitnessis improvedby reducingthe fit- the costs of the extra aggressionrequiredto defend adnessofothersthroughinhibitingthemfrombreeding,rath- ditional resourcesmay be relativelygreatand are borne er than throughan absolute increase in fitness (Verner only by the inhibitingindividual(s).It is thereforeunlikely to the individualwill 1977).Tree Swallows(Tachycinetabicolor)are obligatory thatthe benefitsofsuperterritoriality hole-nesters,and a singlepair can defend additionalsites exceed the costs. Colgan (1979:605) succinctlysummanot essential for successful reproduction(Harris 1979, rizedthis argumentwith his commentthat "Colloquially, Muldal et al., 1985), thereby preventing conspecifics spitefulnesshas become altruism." Since the potential from using those nest sites. Harris(1979) suggestedthat benefitsofsuperterritoriality to the individualarelow, the the main function of surplus nest site defense in Tree evolution of inhibitorybehaviorin Tree Swallowswould Swallowsis to depressthe reproductivesuccessofpotential requirea verylow cost to the aggressionrequiredto defend breeders.However, Robertsonand Gibbs (1982) found extra nest sites. We provide evidence againsta low cost that Tree Swallowsdo not orienttheiraggressivenest de- of aggression.This evidence includes injuriescaused by fense in relationto the location of surplusnest sites and aggressiveencounters,anda seasonaldeclinein aggressive concluded that Tree Swallows defend a circularradius defenseby residents,which suggestsa trade-offbetween around their nest site, which may encompassadditional time and energyexpended on intraspecificnest defense nest sites. A commentaryby Harris(1985) indicatesthat and otheractivitiessuch as parentalcare. the issue of superterritorialityin Tree Swallows is still Ourstudyis basedon two populationsof TreeSwallows unresolved.Here,we arguethatthe conditionsunderwhich at the Queen's University Biological Station, Chaffey's superterritorialbehavior would evolve do not occur in Lock, 50 km north of Kingston, Ontario, Canada.The Tree Swallows. New Land(NL)populationconsistsof about60 pairsthat breed in nest boxes which are distributedover several hayfields.The NortheastSanctuary(NES)populationhas 1 Received 1 April 1985.Finalacceptance17 September about 35 breedingpairs,with both artificialand natural 1985. nest sites over shallowwater.During 1985 in the NL, we 2 Presentaddress:Division of BiologicalSciences,Unicaught9 birdswithinjuriesthatgenerallyconsistedof large versity of Michigan,Ann Arbor,Michigan48109 USA. bare areason the back of the head and neck, and some-