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Predation and Coloniality in Cliff Swallows (Petr 0 Chelid on Pyrrhono Ta )

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Predation and Coloniality in Cliff Swallows (Petr 0 Chelid on Pyrrhono Ta ) PREDATION AND COLONIALITY IN CLIFF SWALLOWS (PETR 0 CHELID ON PYRRHONO TA ) GERALD S. WILKINSON 1 AND GREGORYM. ENGLISH-LOEB2 •Departmentof Biology,University of Californiaat SanDiego, La Jolla, California 92093 USA, and 2Departmentof Biology, San Diego State University, San Diego, California 92182 USA ABSTRACT.--Thehypothesis that the time requiredto detectan approachingpredator variesinversely with colonysize was tested.Ninety-five attacks by LoggerheadShrikes (Laniusludovicianus) and AmericanKestrels (Falco sparverius) were simulatedat six Cliff Swallow(Petrochelidon pyrrhonota) colonies. Colony size, which ranged from 18 to 320nests, did notexplain a significantportion of thevariance in thetime it tookto detectapproaching predatormodels, even when effectsdue to wind velocity,breeding stage, nest exposure, andair temperaturewere removed. Only air temperatureproved to be a significantpredictor of detectiontime. To determinewhether or not large coloniesmight reducethe relative amount of predationmore effectivelythan smallcolonies, we conductedperiodic predator censusesaround each colony and observedactual predator attacks. Relative predator density increasedonly five-fold, while colonysize increased by a factorof 20. Further,we foundno relationshipbetween attack rate and colonysize. Consequently, we suggestthat larger col- oniesmay dilute the effectof localpredators and therebysuffer less nestling predation on a per nest basis.Received 24 April 1981, accepted1 September1981. MOST birds must detect and subsequently attackrate of eachpredator may rise. If nest- evade predatorsif they are to survive. Birds lings are being preyedupon, then the magni- that live in groupsmay have an advantageover tude of the dilution effectwill be greatestwhen solitary individuals, because they can ex- the colonyis highly synchronous.This idea is changeinformation about a predator'slocation quite similar to the predatorswamping or sa- and thereby reduce the risk of predation on tiation hypothesis(Lloyd and Dybas 1966, othergroup members. Although the evolution- Clark and Robertson 1979), which states that ary advantagesof suchalarm calling behavior groupssatiate predators by synchronizingvul- are still in dispute(see Harvey and Greenwood nerabilityin time or space,thus decreasing the 1978,for review), many workersbelieve that probabilitythat any given individualwill be one major evolutionaryadvantage to being in preyedupon. Accordingly,we assessthe va- a largegroup is earlydetection of approaching lidity of the dilution hypothesis by using predators.In this paper we describefield ex- predatorcensuses, direct observationsof at- perimentson the Cliff Swallow(Petrochelidon tempted predation on swallows,and breeding pyrrhonota)that testthe effectof colonysize on synchronyestimates at Cliff Swallowcolonies the time it takesto detectavian predators,and of different sizes. that demonstratethe ability of Cliff Swallows to discrminate predator from nonpredator METHODS models. In addition, we evaluate an alternative Study subject.---CliffSwallows were chosen as advantageto largecolony size, that of diluting subjectsof this investigationbecause they live in an individual's chancesof being captured(Ber- coloniesof differentsizes, emit high-pitched,single- tram 1978). note alarm calls (Samuel1971a), and show a stereo- The dilution hypothesisstates that any in- typeddisplay once an approachingpredator is de- dividual's probabilityof being capturedby an tected.They may build from 15 to severalthousand attackingpredator decreases as the numberof of their gourd-shapedmud nestsbeneath a bridge, overhangingcliff, or building eave (Mayhew 1958, his neighborsincreases, assuming all have an Samuel1971b). Consequently, only avian predators equal chance of being captured. Obviously, cangain access to theirnests easily. American Kes- this dilution effect will always occur unless trels(Falco sparverius) often attack Cliff Swallowcol- predationincreases with colonysize in com- onies(pers. obs.), as theydo BankSwallow (Riparia pensation.This may occur in two ways.Either riparia) colonies(Freer 1973, Windsor and Emlen the number of predatorsmay increase,or the 1975).They attemptto grab perchedadults (Bonnot 459 The Auk 99: 459-467.July 1982 460 WILKINSONAND ENGLISH-LoEB [Auk, Vol. 99 Fig. 1. The four models:American Kestrel, Loggerhead Shrike, Cliff Swallow,and MourningDove. 1921,this study)or flying swallowsor to reachwith- colonyhad ceasedgiving a mobbingdisplay and had in nests for nestlings (this study). Loggerhead resumed normal activity. Shrikes(Lanius ludovicianus) capture most Cliff Swal- We presentedbetween 2 and 12 models(• = 8.4, lowsby pulling nestlingsout of nestswith their bills SE = 0.5) every9 rain (• = 8.9, SE = 0.4) at eachcol- (this study). ony. We beganeach of the 28 daysof presentations Field experiment.--Wesimulated attacks of avian at 0840(SE = 15 min) PacificDaylight SavingsTime. predatorsat six coloniesin San Diego County, Cal- Any trials in which the modelstopped or otherwise ifornia between 6 April and 9 June1978 to determine was towed unusuallywere omittedfrom all analyses. whetheror not largecolonies could detect approach- Excludingthose problemtrials, the mean time, 7.0 ing predatormodels faster than smallercolonies. We s (SE = 0.1, n = 161), required to tow the models toweda stutfedpredator, either a LoggerheadShrike down the guidelines produced a flight speedof 4.3 or American Kestrel, or a stuffed nonpredator con- m/s, which is about one-half our estimates of either trol, either a Cliff Swallow or Mourning Dove (Ze- a shrike or kestrelapproaching a landing. naidamacroura), at the colonies(Fig. 1). The models Using our recordings,written observations,and emergedfrom a smallblind, traveledalong 30 m of the photographs,we measuredthe responsetime horizontalguide lines, which were orientedperpen- and the intensity of the display for each trial. We dicular to the colony stTucture,and then entered usedthe elapsedtime betweenwhen the modelleft another small blind beneath the center of the colony the blind and when the first alarm call was given as (Fig. 2). We designedthe apparatusto mimic an ac- the responsetime. We scoreddisplay intensity on a tual attack; kestrels and shrikes, however, sometimes six-point scale:zero equals no detectableresponse approacheda colonyfrom a greaterdistance or more laterallyto the colonyface. Due to differencesin to- pographyand the nestingstTuctures, the geometry of the apparatuswas similar but unique at eachcol- ony (Table 1). We timed and tape-recordedthe responseusing a stopwatch,Nagra IV-D tape recorderat 7¾2ips, and a SennheiserMKH 815T directional microphone and took a photographof the colonyat the end of the trial from within a blind adjacentto the starting pole. Written descriptionsof the swallows'response were made after each trial, based on a mutual deci- sion by the observers.We randomizedthe order in whichwe presentedthe models(except when testing for habituation and sensitization, in which case we presented the same model for many consecutive Fig. 2. Avian predatorattack simulation appa- trials). The next model was not presenteduntil the ratus. JULY1982] Predationon Cliff Swallows 461 TABLE1. Summary of colony characteristics. Deviation from Deviation Distance Number of Direction perpen- from below (-) or Exposure active nests model dicular to horizontal above (+) (percentage Name of Nesting flown structure approach nests nests colony 1978 1979 structure from (degrees)(degrees) b (m) facing out) EC 320 674 Bridge W 0 0 -2.5 90-95 DM 147 208 Bridge W 35 0 -5.0 85-90 SF 96 100a Bridge E 19 -5 -3.5 0 SP 92 100a Building N 0 +8 -1.5 50 SV 76 92 Bridge E 8 -2 +1.0 0 SM 18 35 Bridge E 0 -2 -1.0 50 Theseare estimatesbased on a winter inventoryof the nestingsite. Negativevalues refer to downwardslope, positive values to upwardslope. and five indicates that the response was indistin- hatchingwas usedas a measureof colonysynchrony guishablefrom that given to real predatorsduring (Emlen and Demong 1975). an actual attack. We based intermediate intensity Predator censuses.--We periodically censused scoreson the proportion of birds in the colony that American Kestrelsand LoggerheadShrikes in a 1- gave alarm calls and entered the mobbing flock. km circlearound each colony during the 1979breed- Alarm callsalways preceded the formationof a mob- ing season. Five observationstations were estab- bing flock. In an actualattack, once an alarm callwas lished within each censusarea, one placed at the given, most birds vacatedtheir nestsand formed a colonyand the othersdistributed a mean distanceof coordinatedflock that wheeledat the predatorwhile 0.6 km (SE = 0.1) aroundthe colony.At eachstation individuals continually gave alarm calls. We com- an observerspent 5-15 min scanningthe surround- paredmedian mobbing response intensities for each ing vegetationfor perching kestrelsor shrikes. Be- model to determine whether or not swallows could causeboth AmericanKestrels (Balgooyen 1976) and discriminate between models. LoggerheadShrikes (Craig 1978)typically sit on el- We measured exposure of the nest, air tempera- evated perches, they are readily observedover long ture, wind velocity, and breedingstage, in addition distances.At most of the colonies,the surrounding to colonysize, for eachtrial, becauseeach may have terrain was primarily level and covered with low causalrelationships with responsetime. If the first vegetation.Both predatorspecies nested within the swallowto give the alarmtends to be sitting in the census area
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