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Interspecific Competition and Social Behavior in Violet-Green Swallows

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Interspecific Competition and Social Behavior in Violet-Green Swallows 606 ShortCommunications [Auk, Vol. 107 Interspecific Competition and Social Behavior in Violet-green Swallows JEFFREYD. BRAWN' Departmentof BiologicalSciences, Box 5640, Northern Arizona University, Flagstaff,Arizona 86011 USA Resourcecompetition can lead to aggressiveor ter- swallowaggression toward female Western Bluebirds ritorial behavioramong conspecifics or heterospecif- aswell (e.g. Prescott1982). Encounters commonly in- ics(Orians and Willson 1964).The demographiccon- volved one or more swallowsflying towards,but not sequencesof these interactions may also influence striking, a bluebird perchedon or near a box. Blue- other life-history traits(Thornhill 1987).Competition birds remained perched, made frequent "rushing" for nest sitesamong cavity-nestingbirds is a potent flightsat oncomingswallows, or left the vicinity. In selective force that often prevents sexually mature 9 instances, as swallows "harassed" a bluebird, other individuals from breeding (Morse 1980). In popula- swallowsentered disputedboxes and depositednest tions of cavity nestersthat breed in north-centralAr- material(pine needles)or removedbluebird nestma- izona (see Brawn and Balda 1988), I observed inter- terial (grass).I never observedswallow aggression specificinteractions between Violet-green Swallows toward bluebirds away from nest boxes. (Tachycinetathallasina) and Western Bluebirds (Sialia Swallows were not marked individually, but I es- mexicana),whose abundancesare limited by nest-site timated groups to range from 4 to 12 individuals. availability.Here I describethese encounters and dis- Thesegroups always included two or more brightly cussthe possibleinfluence of interspecificresource colored males and two or more dully colored indi- competitionon socialbehavior in Violet-green Swal- viduals(probably females, but the phenologyof swal- lows. low plumagecharacteristics is not reliablyknown [Pyle In 1980, I installed 60 nest boxes on each of two et aL 1987]).Solitary swallows were all brightly col- 8.5-haareas in ponderosapine (Pinusponderosa) forest ored males. habitat where natural cavities were scarce(Brawn and Solitary swallowsnever displacedbluebirds (n = Balda1988). These areas are locatedsouth of Flagstaff, 15), but groupsdrove bluebirdsaway from a box in Arizona, in the U.S. Forest Service Coconino National 61% of the observedinteractions (n = 59). The pro- Forest, Coconino County. Swallows and bluebirds portionof groupinteractions that displacedbluebirds within each area used nest boxes almost exclusively was significantlygreater than that for bluebird vs. throughout the study (1981-1984, see Brawn 1988). solitaryswallow encounters(Chi-square test for pro- Neither speciesis a permanentresident, but bluebirds portions, X2 = 24.2, df = 1, P < 0.001). Of the 33 arrive and begin to nestabout a month beforeswal- disputednest boxes, 11 (33%)were usedby swallows, lows. bluebirds remained and nested in 18 (55%), and 4 I observedinterspecific interactions along transects (12%)were not subsequentlyused by either species. (5 per area) that permitted close observation of all One memberof sixdisplaced bluebird pairs was color- boxesduring each visit to an area.These observations banded,and two of thesepairs nested in other boxes. were made during the morning (0530-1100)or late The rate of interspecificinteractions per hour of afternoon(1500-1830) from early April through mid- field observationincreased from 1981through 1984. June. Upon detecting an interspecificencounter, I This trend was concomitant with annual increases in observeda disputed nest box until only one species numbersof boxesoccupied by bluebirdswhen swal- was present for longer than 15 min. Beginning in lows arrived at the study areas (Spearmanrank test, early May, I periodically inspectedall nest boxesto p = 0.64, df = 6, P < 0.10, samplesize based on number determinenesting activity. Boxes with nestswere in- of breedingseasons per areawhen both specieswere spectedtwice weekly throughoutthe nestingcycle. present).At least40% of the boxeson eacharea were I observed 74 interactions between swallows and unusedeach year, but thesewere probablyunsuitable bluebirds at 33 different nest boxes. Bluebirds had for swallowsbecause they were either too low or in begunnesting in all the disputedboxes and therefore densevegetation (see Brawn 1985). A relationship alwayshad priority. Fifty-nine of the encounters(80%) between variation in availability of contestablere- involved a single adult male bluebird and four or sourcesand frequencyof interspecificinteractions has more swallows; the remainder were between a male been reported in other cavity nesters(Minot and Per- bluebird and one swallow. Others have observed rins1986), among nectarivorous birds (Stiles and Wolf 1970),and among scorpionflies(Thornhill 1987). Within a breeding season,swallow aggressionwas ' Presentaddress: Smithsonian Tropical Research commonlydirected at bluebird pairs in the early or Institute,Box 2072, Balboa, Republic of Panama.Cor- preincubationstages of the breeding cycle. On av- respondenceaddress: Smithsonian Tropical Research erage,30% of the bluebirdnests were moreadvanced Institute, APO, Miami 34002-0011 USA. when swallowsbegan to selectnest sites.I never ob- July1990] ShortCommunications 607 served attemptsto usurp these boxes(but seePrescott Slagsvoid1978) would allow experimentalevaluation 1982).Field experimentswith other specieshave also of the relative importance of intra- vs. interspecific demonstrateda positive associationbetween pheno- resourcecompetition in avian social behavior. logical overlap in breeding activity and frequencyof I thank D. Enstrom,K. Johnson,B. G. Murray Jr., interspecificinteractions at nestsites (Slagsvoid 1978, T. D. Pitts, S. K. Robinson, K. Sieving, and B. Stutch- Gowaty 1981). Gwinner et at. (1987) suggesteda prox- bury for constructivecomments on earlier drafts. R. imate explanation; simple physical associationof a P. Baldaprovided usefuladvice throughout the study. male cavity nesterwith a nest site can increasetestes sizeand plasmalevels of luteinizing hormonewhich, LITERATURE CITED in birds, is associatedwith aggressiveor territorial behavior.The magnitudeof this effect increasesas an BRAWN,J.D. 1985. Population biology, community individual is associated with a nest site. Thus, swallow structure, and habitat selection of birds in pon- aggressiontowards bluebirds may be moresuccessful derosa pine forest habitat. Ph.D. dissertation, againstrelatively late breedersthat offer lower re- Flagstaff,Northern Arizona Univ. sistence. 1988. Selectivity and ecological conse- I suggestthat socialbehavior in Violet-green Swal- quencesof cavity nestersusing natural vs. arti- lows enhancesaccess to a vital resource.Interspecific ficial nest sites. Auk 105: 789-791. dominanceis often determinedby body size (Murray --, & R. P. BALDA. 1988. Population biology of 1981)and the averagemass of male Violet-green Swat- cavitynesters in northern Arizona:Do nestsites lows (œ= 14 g) is approximatelyhalf that of male limit breeding densities?Condor 90: 61-71. Western Bluebirds (œ= 29; Dunning 1984). Forming DVNNING,J. B., JR. 1984. Bodyweights of 686 species groupsto overcomeresource defense by larger-sized of North American birds. West. Bird Band. Assoc. heterospecificshas been observedamong other bird Monogr. 1: 1-37. speciesthat competefor accessto food (Murray 1981) GOWAT¾,P.A. 1981. Aggressionof breeding Eastern and amongcoral-reef fish (Robertsonet al. 1976).Ap- Bluebirds (Sialia sialis) toward their mates and proximately25% of all swallow nests(n = 47) were modelsof intra- and interspecificintruders. Anita. in boxespreviously occupied by bluebirds. Thus, so- Behav. 29: 1013-1027. cial aggressiontowards heterospecificsis not man- GWINNER,H., E. GWINNER,& J. DITTAMI. 1987. Effects datory to obtain a nest site. Nonetheless,interspecific of nestboxes on LH, testosterone, testicular size, competition(exploitative and interference)appears and the reproductive behaviour of male Euro- particularlystrong for swallowsin north-centralAr- pean Starlingsin spring. Behaviour 103: 68-82. izona becausethey arrive at breeding habitat and LOMBARDO, M. 1986. Attendants at Tree Swallow commence reproduction after all other sympatric nests.I. Are attendantshelpers at the nest?Con- speciesof cavity nesters(n = 5; see Brawn and Balda dor 88: 297-303. 1988). MINOT, E. O., & C. M. PERRINS.1986. Interspecific I frequently observed2 or more individuals enter interference competition-nestsites for Blue and active swallow nests, but I could not ascertain wheth- Great tits. J. Anita. Ecol. 55: 331-350. er or not all birds actually assistedthe nest attempt. MORSE,D. H. 1980. Behavioral mechanisms in ecol- The question of whether the swallow groups that ogy. Cambridge, Massachusetts,Harvard Univ. engaged in aggressiveinteractions were temporary Press. aggregationsor persistentsocial units is unanswered. MURRAY,B. G., JR. 1981. The origins of adaptive If Violet-green Swallows are true cooperativebreed- interspecificterritorialism. Biol. Rev. 56: 1-22. ers, my observationsmay indicate a rare example of OR•^NS,G. H., & M. F. WILLSON.1964. Interspecific interspecificresource competition that affected the territories in birds. Ecology45: 736-745. evolution of a resource-based social system (see ROBERTSON,D. R., H. P. A. SWEATMAN,E. A. FLETCHER, Thornhill 1987). Alternatively, Tree Swallows (T. bi- •r M. G. CLELAND.1976. Schooling as mecha- color)at active nests include a breeding pair and un- nism for circumventing the territoriality
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