Nest Material Preferences of Great Frigatebirds
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j. Field Ornithol., 56(3):236-245 NEST MATERIAL PREFERENCES OF GREAT FRIGATEBIRDS BY LINCOLN FAIRCHILD, SHEILA A. MAHONEY, AND RALPH W. SCHREIBER Analysesof variouspasserine nests (Kilgore and Knudsen 1977, Collias and Victoria 1978, Jacobset al. 1978, Thomas 1983) and laboratory studies(Sargent 1965) have shownsome nesting material preferences. No suchstudies have been done on marine species,and none has com- pared selectednesting material to material availabilityin the environ- ment. We comparedthe nesting material in nestsof Great Frigatebirds (Fregataminor) on EasternIsland, Midway Atoll, PacificOcean, to that availablein the environment.We experimentallytested whether Great Frigatebirdsprefer certain typesof sticksor merely take sticksin pro- portion to their availability. METHODS We conductedour study during May and early June 1983, in the frigatebirdcolony located on the windward,south central side of Eastern Island, Midway Atoll which contained76 nestsbuilt in the crownsof Scaevolabushes about 2 m above the ground. Inter-nest distancesin the sameclump were highly variable(approximate range 1 to 10 m). At the beginningof the study,most nesting frigatebirds were incubating(n = 59), but 5 pairs brooded chicks;2 pairs had just formed and had not producedeggs; and 10 maleswere still displaying.By the end of the study,most nests still containedeggs (n = 36), but 22 nestscontained chicks,and no maleswere displaying.Thirteen nestshad been aban- doned. We examined 6 recently abandonednests. Five of the 6 nestswere examinedimmediately after abandonmentprior to any nestdestruction by other frigatebirds.The sixthnest was examined several hours after abandonmentbut appeared intact. We measuredthe length of each stickin each nestto the nearestmillimeter and quantifiedthe sizeand numberof branchesof all sticksby assigninga valueof 1.0 to a branch if the branch was about the same diameter as the main stem of the stick. We assigneda value of .5 to a branch if the branch diameter wasless than the stem diameter. The sum of all values of branches of a stick is the BranchingIndex (BI) of the stick. We log transformedall stick lengthsand branchingvalues. Since the BI of a stick containingno brancheswould be zero and would not allow usto log transformbranch- ing data, we added one to all BI's prior to log transformation.Thus, the BI of a stick without branchesequalled 1.0. If BI -- 2.0, the stick contained either one large branch or two small branches.We used ANOVA to test whether differencesin length or branching existed between nests. 236 Vol.•6, •o. • FrigatebirdNests [237 We surveyedthe ground around the nestingarea for sitesthat con- tained sticks.Open areasthat were smallerin any directionthan about 2 m (the wing spanof frigatebirds)were not consideredbecause frigate- birdsprobably could not easilyland or hover and henceretrieve sticks from them. We sampledsticks on 5 main pathsand 4 fieldsin the nesting area. We deliberatelyselected, as one of the four fields,a sitewith many large sticksthat appearedappropriate for frigatebird nests. We useda m2 quadrat to sampleground sticksand adopteda sampling regime to determine whether the proximity of the sampleto the edge of the path or field affectedstick distribution. Sticks were sampledevery 10 paces,or approximately6.6 m, along each path: the first sampleon the right side of the path, the secondsample in the path center, and the third sampleon the left sideof the path. This sequenceof sampling wasrepeated along the length of the path and a similarstaggered sam- pling regimewas used for fields:we took samplesalong both sides,along the field'smidline, and half way betweenthe midline and either sideof the field.At eachsample location, we measured and assigned a branching index to all sticksthat were at least 10 cm long and totally or partially withinthe m2 quadrat. Stick lengths and branching were log transformed and analyzedby ANOVA to test whether differencesexisted among quadratswithin a given area. Differencesamong quadrats were found only in the area we deliberatelyselected because of the high densityof large sticks.Excluding this area, quadratswere pooledwithin areasand ANOVA then testedwhether differencesexisted among areas. To test preferencesin sticks,we asked two questions:(1) Is stick positionwith respectto the ground important to frigatebirds?And (2) doesstick type and/or positionwith respectto the groundaffect frigate- bird stickselection? To answerthe first question,we collectedstraight sticksof variouslengths and dividedthem into 2 groupsequal in number and averagelengths. We painted the tips and middle of the sticksby group for later identification.To control for paint color, the two stick groupsdiffered only in the sequenceof red and blue bandsof paint. We placedall sticksfrom one groupsingly on the topsof Scaevolabushes near the nestingcolony. Sticks from the other groupwere placedsingly on the ground at the edge of a field in front of the samebushes. In the first of two trials, we used32 sticksin each group and left them unat- tendedafter placement.Two dayslater, wesearched the areathoroughly and collectedand countedall remainingsticks. We surveyedfrigatebird nestsand the ground around the nestingarea daily for painted sticks. Our cautiousnest surveys had only a minimal effect on nestingbirds at the possibleexpense of failingto sightpainted sticks that mayhave been incorporatedinto nests.In the secondtrial, we used20 sticksper group and observedall activitynear the bushand groundcontaining our sticks for one hour followinginitial placement. To answerthe secondquestion, in each of 12 trials, we constructed two nestseach with 20 branchedsticks and placedone in a Scaevolabush and the other on the ground in front of the bush. We constructeda 238] L. Fairchitdet at. j. FieldOrnithol. Summer 1985 T^BI•E1. Dimensionsof stickson the groundin pathsand fieldsand in frigatebirdnests. The mean density,length, and branchingindex ___1 standarddeviation are presented. n Area (quad- n Density Length Branching Site (m2) rats) (sticks) (sticks/m2) (cm) index Path 1 473 15 135 9.0 __+8.5 21.6 __+11.7 1.08 __+1.24 Path 2 466 9 70 7.8 ___5.7 18.6 ___9.8 1.04 ñ 1.19 Path 3 299 15 118 7.9 __+7.8 20.4 ñ 14.2 1.11 __+1.30 Path 4 1090 15 144 9.6 __+6.5 21.1 __+18.4 1.09 __+1.28 Path 5 301 15 148 9.9 __+12.7 18.8 __+11.5 1.07 __+1.25 Field 1 1026 15 234 15.6 ñ 13.8 19.1 ñ 12.5 1.08 __+1.24 Field 2 334 10 95 9.5 _ 11.5 21.3 ñ 10.1 1.06 ñ 1.23 Field 3 452 14 209 14.9 __+12.8 18.2 __+8.3 1.07 __+1.30 Field 4 78 5 271 54.2 __+43.0 21.5 __+14.2 1.07 ñ 1.21 Nest 1 30 41.8 __+13.0 1.98 __+1.72 Nest 2 30 44.9 __+21.7 2.20 --+ 2.06 Nest 3 47 42.5 __+14.5 1.99 ñ 1.88 Nest 4 35 45.9 __+15.3 1.64 __+1.53 Nest 5 48 36.8 __+16.1 1.86 __+1.70 Nest 6 46 39.3 __+15.4 1.90 ñ 1.50 All sites(ground) 4510 113 1424 20.0 --+12.0 1.08 __+1.25 All nests 236 41.4 __+16.1 1.90 __+1.80 third nestof 20 stickscontaining no branchesand placedit on the bush within 2 m of the branchedstick nest. To control for possibledifferences in our selectednest locations, we reversedthe positionof the two nests in the bush in each experiment. All nestswere constructedfrom sticks obtained from abandonedfrigatebird neststo diminish the possibility of using sticksthat were inappropriatefor frigatebirds.We observed thesenests from about 75 m awayfor 1 h and recordedthe numberof landingson eachnest, the cumulativetime birds spenton each nest,the number of attemptsby birds to remove sticksfrom each nest, and the number of sticksactually removed and carried from each nest. At the end of the hour we disassembled all nests and counted the number of sticksremaining. The above experiment confoundsstick type and apparent nest size. Althoughall nestscontained 20 sticks,branched sticks are more massive than stickswithout branches.Thus, the branched stick nestsappeared more massivethan the straightstick nests. To control for the effect of the apparent nest size in our third experiment, we constructeda small nest with 20 branched sticksand a large nest with 30 stickscontaining no branches.We placedboth nestsin a Scaevolabush within 2 m of each other. About 18 h later, we disassembled the nests and counted the numberof remainingsticks in each. RESULTS Frigatebirdnests averaged 39.3 ___8.6 sticks(n -- 6 nests).Stick density in all pathswas 8.5 ___8.3 sticks/m2 (n = 69 quadrats),and no significant Vol.56, No. 3 FrigatebirdNests [2 39 45 GROUNDN -- 1424STICKS 40 35 NESTN -' STICKS236 30 25 20 15 10 5 10 15 25 30 35 5 70 75 80 85 86+ STICK LENGTH (cm) FIGURE1. Groundand neststick lengths. The solidbars represent ground sticks, and the hatchedbars represent nest sticks. Sticks are lumpedinto 5-cm sizeclasses for illustrativepurposes only. differenceexisted among the 5 paths(F,64 = .17, P > .05, Table 1). While therewas no significantdifference in stickdensity in 3 of the 4 fields(F2.•4 = 1.94,P > .05),the density in thesefields was significantly greaterthan in paths(F,104 = 8.76, P < .005). The averagedensity in fieldswas 13.8 _ 12.8 sticks/m2 (n = 39 quadrats).The averagestick densitywas greater in the fourth field, 54.2 _ 43.0 sticks/m2 (n = 5 quadrats). Averagestick length in nestswas 41.4 _ 16.1 cm (n = 236), whereas the averagelength of groundsticks was only 20.0 _ 12.0cm (n = 1424, Table l) and significantlyshorter than neststicks (F1,1658 = 628.1, P < .001, seeFig. 1). The differencebetween the lengthof groundand nest stickswould be evengreater had we includedin our groundsamples the numeroussticks that were lessthan 10 cm long.