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Significance of the Pattern of Nest Distribution in the Pigeon Guillemot (Cepphus Col Umba )

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Significance of the Pattern of Nest Distribution in the Pigeon Guillemot (Cepphus Col Umba ) SIGNIFICANCE OF THE PATTERN OF NEST DISTRIBUTION IN THE PIGEON GUILLEMOT (CEPPHUS COL UMBA ) S. K. EMMS • AND N. A.M. VERBEEK Departmentof BiologicalSciences, Simon Fraser University, Burnaby, British Columbia V5A 1S6,Canada ABSTRACT.--Westudied nest distribution in the Pigeon Guillemot (Cepphuscolumba), and three possibleselective factors that influence it: nest predation, the use of nest groupsas information centers for food finding, and spatial variation in nest-hole quality. Quadrat surveysin 2 yr suggestedthat nestswere dispersedrandomly with respectto the distribution of suitableholes, but on a larger spatialscale, holes on the south shorewere preferred to those on the north. Predation intensity was largely independent of nest density. This was true whether egg predationalone, chickpredation alone, or total predationwas considered. Nestingclose to other birds, however,had a slight advantageon a very local scale.Reduced predationon south-shorenests seemed to be due to their beingbetter protected from predators ratherthan to specificadvantages to aggregation.Neither chickfeeding rates nor chickgrowth ratesincreased with nestgroup size, indicatingthat nestgroups did not serveas information centersfor food finding. We concludethat the observedclumping of nestswas due to a clumpeddistribution of suitablesites and spatialvariation in nest-holequality. Received18 February1988, accepted3 October1988. COLONI^Lnesting is a highly conspicuous knowledgeableroost mates, and recentlyBrown pattern of breedingin birds, and many studies (1986), Greene (1987), and Waltz (1987) provid- havebeen devotedto understandingits adap- ed strong evidence that Cliff Swallow (Hirundo tive significance(see Wittenberger and Hunt pyrrhonota), Osprey (Pandion haliaetus), and [1985]for review). Lack(1968), in a majorcom- Common Tern (Sternahirundo) colonies serve as parative study, found that coloniality was dis- information centers. proportionately common in speciesthat eat A secondmajor explanation for the evolution aquaticprey, seeds,or fruit, and that it is often of coloniality is that predation pressure de- associatedwith flockfeeding. He suggestedthat creasesas nest density increases(Patterson 1965, the main advantageof coloniality, at least for Robertson 1973, Parsons 1976, Hoogland and species that feed in flocks, was an increased Sherman 1976, Birkhead 1977, G0tmark and An- efficiencyof food exploitation. Ward and Za- dersson 1984, Robinson 1985). Several mecha- havi (1973) argued explicitly that many bird nisms have been invoked to explain this fact, speciesused breeding colonies as "information although alternativesare often difficult to sep- centres"for food finding. Severalstudies have arate in the field. First, predator mobbing often supportedWard and Zahavi'shypothesis (Horn increaseswith colony size or nesting density 1968, Siegfried 1971, Krebs 1974, Emlen and (Patterson 1965, Tenaza 1971, Balda and Bate- Demong 1975, Erwin 1978, Loman and Tamm man 1972, Hoogland and Sherman 1976, GOt- 1980, Ydenberg et al. 1983), but Bayer (1982) mark and Andersson 1984, Robinson 1985). In- arguedthat other hypotheses explained equally creasedmobbing may drive off predators or well much of the data, and Andersson et al. lower their hunting success.Second, the per in- (1981) found that Black-headedGull (Larusri- dividualprobability of being caught decreases dibundus)colonies did not appear to serve as ascolony size increasesthrough a prey dilution information centers.However, De Groot (1980) effect (Bertram 1978). Third, a "selfish herd" showedunequivocally that captiveQuelea que- effect may operate (Hamilton 1971). Fourth, lea learned the location of food or water from predator satiation may occur if prey are suffi- ciently abundant. Support for this last mecha- nism comes from studies that show that the • Present address:Department of Biology, Prince- proportion of individuals killed decreasesdur- ton University, Princeton, New Jersey 08544-1003 ing the peak of the breeding season(Patterson USA. 1965, Nisbet 1975, Daan and Tinbergen 1979, 193 The Auk 106:193-202. April 1989 194 EMMS^•qD VERSEEK [Auk,Vol. 106 Findlay and Cooke 1982, Nisbet and Welton The north and east shores are much lower and more 1984). gently sloping. Exfoliation of the rocks is most ex- A third possibleinfluence on the pattern of tensive on the east and south shores, where there are a number of boulder tumbles below the cliffs, and nestdistribution is spatialvariation in siteavail- here the majorityof the guillemotsbreed. Details about ability. Birds may nest in clumpsbecause suit- the islandare given in Butler (1974, 1980)and Brooke able sitesare limited and patchily distributed et al. (1983). Thomson (1981) provides detailed in- (Lack 1968, Snapp 1976), or becausehigh-qual- formation about the oceanographyof the Strait of ity sitesare clumpedeven though suitablesites Georgia. are not limited. Approximately200 pairsof PigeonGuillemots nest Most studiesof colonialityhave concentrated on Mitlenatch. We determined their breeding distri- on highly colonialspecies (e.g. Patterson1965, bution by systematicallysearching the island shore Tenaza 1971, Parsons1976, Hoogland and Sher- for nestsand mapping their locationsonto a 50 m2 man 1976, Birkhead 1977). Loosely colonial quadrat system.(We did not searchthe cliffs on the species(those that nest in smaller, less dense south shore to avoid disturbing breeding cormo- rants.)We found nestsduring the egg-layingperiod, aggregations)have received lessattention, but or occasionallylater in the season,by observingfood the factors that influence their nest-distribution deliveries to chicks.We searchedall quadratsfor un- patternsmay be very different.Studies of loose- occupiedbut apparentlysuitable nest holes. We mea- ly colonial speciesmay also help to illuminate suredthe height and width of the entrancehole, the the problem of the evolutionof coloniality,be- distancefrom the hole entranceto the nest,the height causetheir nest-distribution patterns are prob- and width of the nest chamber, and the distance from ably similar to those on which selectiononce the hole entrance to the rear of the chamber. We acted in speciesthat are now highly colonial. scored concealment of the hole entrance on a scale Thusone canpartially avoid the argumentthat of one to five, and complexity of the nest chamber on a scale of one to six. Holes were considered suitable by studying highly colonial speciesone can if the measurementsfell within the range of the same demonstrateonly the selectivefactors that main- measurementsof occupiedholes. Birds might have tain coloniality. We studied the Pigeon Guille- used other criteria to decide whether a hole was suit- mot (Cepphuscolumba) because it nestsat a wide able or not. Although our criteria might have led to range of densities, from solitary individuals to an under- or overestimation of the total number of loose aggregationsand even dense colonies in suitableholes, it is unlikely that we createda system- somehigh arcticregions (Nettleship and Evans atic bias in the data. 1985).Our purposewas to determinethe factors To look for nonrandompatterns of nest distribu- responsiblefor the nest-distributionpattern of tion, we plotted nestdensity per quadratagainst suit- this species. able-holedensity and comparedthe fitted slopeto the slopeexpected if nestswere distributedrandomly. A random distribution of nestsimplies that the same STUDY AREA AND METHODS proportion of suitableholes is occupiedat all hole densities,so the expectedslope has a value equal to The Pigeon Guillemot is a medium-sized(400 g) the overallproportion of occupiedholes. If birdschose alcid that breeds on the coasts of the northern Pacific. to nest colonially, proportionatelymore holes than It nests in holes under boulders and rock tumbles on expectedwould be occupiedat high hole densities the lower shore, in crevices in rock faces, in driftwood and proportionatelyfewer holesthan expectedwould piles, and in a variety of similar locations(Drent et be occupiedat low hole densities.The fitted slope al. 1964),including man-madesites (Campbell 1977). would then be steeperthan expected.Conversely, if It is unusualamong the alcidsbecause it nestslargely birdswere territorial, the fitted slopewould be shal- in small,loose aggregations, feeds mainly on inshore, lower than expected.The data were transformedbe- benthic organisms,and lays normally two eggsin- fore analysisbecause nest density per quadratis bi- stead of one (Storer 1952, Thoreson and Booth 1958, nomially distributed (a hole is either occupiedor Drent 1965, Koelink 1972, Kuletz 1983). unoccupied).Its variance therefore increasesas suit- The study was done on Mitlenatch Island, British able hole density increases.Homogeneity of vari- Columbia(49ø57'N, 125ø00'W), a small(35.5 ha) rocky ances was obtained by dividing both sides of the islandat the northernend of the Strait of Georgia,in regressionequation by the squareroot of the inde- 1984(between 20 April and 24 August) and 1985(be- pendentvariable: the regressionwas transformed from tween 20 April and 18 August). The island is com- N = bSto N/V'• = bV'•, whereN = nestsper quadrat posedlargely of basalticrock that formstwo low hills and S = suitable holes per quadrat. Becauseb is the (<60 m) and a seriesof rocky bluffs. The south shore samein both cases,the slopefitted to the transformed consistsof low, vertical cliffs up to 20 m in height. data can still be compareddirectly to the expected April1989] PigeonGuillemot Nesting Dispersion 195 slope.One potential
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