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Plumage Wettability of Aquatic Birds January1984] ShortCommunications 181 TABLE 1. Latitude and elevation of sites where field studies of Bobolinks have been conducted. Lafi- Eleva- rude tion Location (øN) (m) Source DouglasLake, CheboyganCounty, Michigan 45033' 219 O. S. Pettingill, Jr. (pets. comm.) Sauk City, Dane County, Wisconsin 43016' 231 Martin (1971) Lakeport,Madison County, New York 43010' 113 R. L. Kalinoski (pers. comm.) Bridgeport,Madison County, New York (ShackeltonPoint) 43010' 114 This study P-Ranch,Harney County, Oregon 42049' 1,400 Wittenberger (1978) Brooktondale, Tompkins County, New York (Bald Hill) 42021' 490 This study Adult Bobolinksroutinely left their territoriesto for- COPY,M. L. 1971. Ecologicalaspects of reproduc- age in patchesof winter cressand were observedto tion. Pp. 461-512 in Avian biology, vol. 1 (D. S. carry 1-3 cabbagebutterfly larvae to nestsknown to Farner and J. R. King, Eds.).New York, Academ- contain nestlings. Lepidopteran larvae were an im- ic Press. portant food for Bobolinknestlings in Oregon (Wit- ENGELS,W.L. 1969. Photoperiodicallyinduced tes- tenberger1980). It may be that the density and dis- ficular recrudescencein the transequatorial mi- persionof the hostplant, in combinationwith spring grant Dolichonyxrelative to natural photoperi- weatherfavorable to first generationcabbage butter- ods. Biol. Bull. 137: 256-264. flies, produceda unique situation locally, allowing KLUIJVER,H.N. 1951. The population ecologyof the double-broodedness in Bobolinks. great tit, Parusm. major.Ardea 39: 1-135. I greatly appreciatethe assistanceof G. H. Farley, LACK,D. 1968. Ecologicaladaptations for breeding S.C. Moore, T. J. Rawinski, and D. A. Takacs in the in birds. London, Chapman and Hall. field. Temporal continuity of the project could not MARTIN,S.G. 1971. Polygyny in the Bobolink:hab- have been maintainedwithout the cooperationof J. itat quality and the adaptive complex. Unpub- L. Forney, Director of the Cornell Biological Field lished Ph.D. dissertation.Corvallis, Oregon, Or- Station, and F. Liddington, owner of the Bald Hill egon State Univ. hayfields.D. K. Dawson,B. G. Murray, Jr., V. Nolan PERRINS,C. g. 1965. Population fluctuations and Jr., R. T. Reynolds,P. W. Sherman, C. R. Smith, and clutch-sizein the great tit, Parusmajor. J. Anim. J. F. Wittenberger provided pertinent commentson Ecol., 34(3): 601-647. the manuscript.This researchwas supportedby the WITTENBERGER,J. F. 1978. The breeding biology of Cornell University Agricultural Experiment Station, an isolated Bobolink population in Oregon. Con- New York State College of Agriculture and Life Sci- dor 80: 355-371. ences. --. 1980. Vegetation structure,food supply, and polygyny in Bobolinks (Dolichonyxoryzivorus). LITERATURE CITED Ecology61: 140-150. BENT, A.C. 1958. Life histories of North American Received14 February1983, accepted8 luly 1983. blackbirds,orioles, tanagers and allies. U.S. Natl. Mus. Bull. 211. Plumage Wettability of Aquatic Birds SHEILA A. MAHONEY Departmentof BiologicalSciences, Florida Atlantic University, Boca Raton, Florida 33431 USA Rijke (1970) investigatedthe feather structureand water penetration through the feather layer. Body wettability of breastfeathers of 32 aquaticand ter- feathersand flight featherswould be expectedto have restrial bird families. He found that the breast feath- different characteristics, however, as a result of dif- ers of terrestrial families tended to be more water ferencesin structure,feather density, and packing; repellanton the surface,whereas the feathersof fully furthermore, maintenance activities, such as preen- aquaticfamilies tended to have greater resistanceto ing, also affect wettability. 182 ShortCommunications [Auk,Vol. 101 iO 9 8 7 6 RP .TBp 1(3) 5' (5) 4 3 2 I .EG .•DCC (•) (5) 0 500 I000 1500 2000 2500 30 0 BODYWEIGHT (g) Fig. 1. Plumagewettability of selectedaquatic birds. RP = Red Phalarope;ST = SootyTern; EG = Eared Grebe; CG = California Gull; WG = Western Gull; A = Anhinga; DCC = Double-crestedCormorant; GF = Great Frigatebird;BP = Brown Pelican.Means are shown plus or minus one standarddeviation (vertical lines). Numbers in parenthesesare sample sizes. I define plumage wettability as the percentagegain Bird Care Center in Fort Lauderdale, Florida kindly in body weight that a wet bird experiences.Because allowed me to sample captive Brown Pelicans(Pele- it is a whole body measurement, it does not allow canusoccidentalis). one to distinguish between water repellancy and Plumage wettability was measuredby obtaining a water penetration, as Rijke's model of individual bird's dry weight and then submerging the bird in feathersdoes. Plumage wettability has important en- water for severalseconds and immediately weighing ergetic consequencesfor aquaticbirds. Wet birds are it again. I held the birds loosely, allowing them to heavier, and they have increasedwing loading and relax their wings while I pushed them completely higher energetic costsin flying. Additionally, heat underwater. Some birds struggled but were unable loss from wet birds is greater, becausethe thermal to open their wings completely. In most casesbirds conductivityof water is 25-30 times that of air. were weighed in a mesh bag, the weight of which Becausethe only data on wettability of the entire was subtractedfrom the total weight. Anhingas and body are anecdotal,I examined plumage wettability cormorantswere placed on a top-loading balanceaf- in a variety of aquaticbird speciesin order to deter- ter a single brisk shaketo remove excesssurface water. mine whether or not any generalpatterns exist among All birds were submergedin fresh water except the aquatic birds. terns and frigatebirds,which were submergedin sea I examinedplumage wettability in nine aquaticbird water. A series of experiments with the California speciesrepresenting seven families; Podicipedidae, Gulls, in which fresh water and Mono Lake water Phalaropodidae,Phalacrocoracidae, Anhingidae, Pel- (2.5 times the concentration of sea water) were used, ecanidae, Fregatidae, and Laridae. All individuals indicated that plumage wettability did not differ in were live, healthy birds. The Double-crested Cor- the different waters. morants(Phalacrocorax auritus) and Anhingas (Anhin- Thermal conductance(K) of Anhingas and cormo- ga anhinga)were captured in Florida and held at the rantswas calculatedfrom a rearrangementof the heat Duke University aviary and were 9-12 months old. balance equation: The Great Frigatebirds(Fregata minor) and SootyTerns (Sternafuscata) were adults caught wild at Midway K = (M - E)/(T•- T•), Atoll, Pacific Ocean. The Eared Grebe (Podicepsnigri- coilis)and California Gulls (Laruscalifornicus) were where M is metabolic rate, E is evaporationrate, T• caughtwild at Mono Lake, California. The gulls were is body temperature (approximatedby cloacal tem- iramatures of 6-8 weeks of age. JosephJehl, Jr. gen- perature), and Tais ambient temperature(for details, erouslysampled the Red Phalarope(Phalaropus fuli- see Mahoney 1981). caria) and Western Gull (Larus occidentalis)at Hubbs- I found no association between body size and SeaWorld, San Diego, California. The S.P.C.A. Wild plumage wettability (Fig. 1), but there was an asso- January1984] ShortCommunications 183 ciation with feeding habits. The six speciesthat are surfacefeeders gained approximately6% when wet. F--1 NormalPlumage Of these,Brown Pelicansalso plunge dive but spend 6 mostof their time floatingon the water surface.Sooty LtJ PoorCondition Terns and Great Frigatebirds are unusual among aquaticbirds, because they do not normally alight on Molted 5 the water but rather pluck prey from the surface. The remainingthree species(Eared Grebe, Double- crestedCormorant, and Anhinga), which forage by diving and swimming underwater for prolonged pe- riods and to variable depths, had lower plumage weftability. Cormorantsand EaredGrebes dive and activelyswim underwaterto seizeprey. Their plum- age wettability was only 1% of their body weight. 3 Anhingas were more wettable, gaining 3% of their body weight. Owre (1967)has suggested that Anhin- gas' greater wettability, as comparedto cormorants, 2 decreasestheir bouyancyand helps them to remain submergedwhile they swim slowly and stalk prey. The condition and amount of feathers also affects wettability. The captive cormorantsin poor condi- I tion gained three times more weight than those in normal plumage(Fig. 2). Anhingasthat had molted primaries and rectrices(lost simultaneouslyin this species)gained two-thirds of the weight that wet an- (6)(3) (8)(5) hingas in normal, full plumage gained (Fig. 2). Therefore, primariesand rectricesaccount for one- DOUBLE-CRESTED ANHINGAS third of the plumage wettability of anhingas. CORMORANTS At least somemembers of all speciesused in this Fig. 2. Plumage wettablility of cormorantsand study could take off in flight when wet. SootyTerns Anhingas.Histograms represent means, and vertical took off with difficulty,but all becameairborn. Of 6 lines indicateplus or minus one standarddeviation. frigatebirds,only 2 (of 4) males could take off from Numbers in parenthesesare samplesizes. the water when wet and then only with considerable difficulty.Frigatebirds have the lowestwing-loading of all seabirds(Murphy 1936). Magnificent Frigate- bird (Fregatamagnificens) males weigh 10% lessthan males (Table 1). Only two of the four experimental femalesand have 11%lower wing-loading (Hatring- males and none of the females were able to take off ton et al. 1972, see also Table 1). Great Frigatebird from the water surfacewhen wet, althoughmost birds
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