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Diving Depths of Shearwaters

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Diving Depths of Shearwaters July2001] ShortCommunications 755 and female age in the primary reproductiveout- Department of Zoology, Uppsala University, put of Pied Flycatchers.Oecologia 126:339-344. Uppsala, Sweden. SHELDON,B.C., AND S. VERHULST.1996. Ecological WIEHN, J., AND E. KORPIM•KI. 1998. Resource levels, immunology: Costly parasite defenses and reproduction and resistanceto haematozoanin- trade-offs in evolutionary ecology. Trends in fections. Proceedingsof the Royal Society of Ecologyand Evolution 11:317-321. London, Series B 265:1197-1201. SIIKAM.,•KI, P., O. RATTI, M. HovI, AND G. F. BENNETT. WIEHN, J., E. KORPIM•KI, AND I. PEN. 1999. Haema- 1997. Association between haematozoan infec- tozoan infections in the Eurasian Kestrel: Effects tions and reproduction in the Pied Flycatcher. FunctionalEcology 11:176-183. of fluctuatingfood supply and experimentalma- STEARNS,S.C. 1992. The Evolution of Life Histories. nipulation of parental effort. Oikos 84:87-98. Oxford University Press,Oxford. WILLIAMS,J. B. 1997. Energeticsof avian incubation. SVENSSON,L. 1992. IdentificationGuide to European Pages 375-415 in Avian Energeticsand Nutri- Passerines.Fingraf AB, Stockholm,Sweden. tional Ecology (C. Carey, Ed.). Chapman and THOMSON, D. L., P. MONAGHAN, AND R. W. FURNESS. Hall, New York. 1998. The demands of incubation and avian clutch size. BiologicalReview 73:293-304. WIDEMO,F. 1989.Effect of blood parasiteson the Col- Received5 August2000, accepted25 January2001. lared FlycatcherFicedula albicollis. Honors thesis, Associate Editor: C. Blem The Auk 118(3):755-759, 2001 Diving Depths of Shearwaters ALAN E. BURGER • BirdLifeSeychelles, P.O. Box 1310, Mont Fleuri, Seychelles ABSTRACT.--Maximumdiving depths were mea- merskirchand Sagar1996, Weimerskirch and Cherel sured for shearwatersbreeding on Cousin Island, 1998, Keitt et al. 2000). More sophisticatedtime- Seychelles.Eighty-three percent of 23 Wedge-tailed depth-recorders(TDRs), which indicatetime spentat Shearwaters (Puffinus pacificus)dived, and their various depths and trace individual dives, have re- mean maximum depth was 14 m (SD = 23 m, range vealed muchof the underwaterforaging behavior of 1-66 m, N = 19). All Audubon's Shearwaters (P. lher- penguins, alcids, and cormorants (Kooyman 1989, minieri)dived, and their mean maximum depth was Croll et al. 1992, Wilson 1995, Watanuki et al. 1996), 15 m (SD = 12 m, range 6-35 m, N = 7). Thesedata but have not been applied to shearwaters.I report contradict the hypothesis that tropical shearwaters maximum depths attained by two tropical shearwa- should not specialize in underwater foraging. They ters, Wedge-tailed(Puffinus pacificus) and Audubon's are capableof exploiting deep prey unavailableto (P. lherminieri)shearwaters, breeding on Cousin Is- most other tropical seabirds.Five Puffinusspecies land, Seychelles(4ø20'S; 55ø40'E), and review data on (temperate and tropical) attained allometrically diving depths of other shearwaters.I examine the scaledmaximum depths comparableto thoseof pen- hypothesisthat tropical shearwatersshould not spe- guins and alcids. cialize in underwater foraging (Brown et al. 1978), It has long been known that shearwatersdive be- and discuss the role of diving in tropical shearwaters. neath the oceanto forage,using their feet and wings for propulsion (Brown et al. 1978, 1981), and show Methods.--Maximum-depth gauges were made anatomical adaptations for this mode of foraging from flexible plastic tubing with internal diameter (Kuroda 1954, Warham 1990). Simple depth gauges 0.8 mm, lined with a thin layer of icing sugar and measuring the maximum depths indicate some re- sealed at one end. Gaugeswere 70-120 mm long on markably deep dives attainedby shearwaters(Wei- Wedge-tailedShearwaters, and 70-90 mm long on Audubon'sShearwaters. As the gaugeis submerged, the increasingpressure forces water into the tube, • Presentaddress: Department of Biology,Univer- dissolving the sugar and leaving a record of the sity of Victoria, Victoria, BritishColumbia V8W 3N5, deepestdive (Kooyman et al. 1971, Burger and Wil- Canada. E-mail: [email protected] son 1988, Hedd et al. 1997). The equation provided 756 ShortCommunications [Auk, Vol. 118 by Burger and Wilson (1988) was used to calculate []Wedge-tailed 1 maximum dive depths. ß Audubon's • Cousin Island supportsbreeding populationsof -13,000 pairs of Wedge-tailed and 5,000-10,000 pairs of Audubon's shearwaters (Burger and Lawrencein press).At the time of this study (7-17 June 1999), Wedge-tailedShearwaters were return- ing nightly to nestcavities for courtship,but did not haveeggs or chicks.Audubon's Shearwaters breed all year round on CousinIsland and the statusof those 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 usedin the study was not known. Adult shearwaters Depth (m) sittingin or nearnest cavities were caughtand band- ed at night. Eachgauge was firmly tied to the baseof FIG. 1. Frequency distribution of maximum a singletail featherwith a thread (dental floss).Once depths attained by Wedge-tailedand Audubon's attached the gaugeswere surrounded by feathers shearwaterson Cousin Island, Seychelles.X-axis la- and createdno impedimentsto the birds. Handling bels indicate the upper limit of each 5 m depth time was 10-15 min. Gaugesnot recoveredwould category. eventuallywork looseor be lostwhen the birdsmolt- ed. Eachbird was sampledonce. I made no attempt to collectfood samples,and none of the captured birds regurgitatedfood. ployment, althoughthe error with dives >20 m was Prolongeddeployment increases errors in these likely to be <10% (Burgerand Wilson 1988).There is devices(Burger and Wilson 1988).I includedgauges also a greaterprobability of recordingan exception- recoveredwithin two daysof deployment,and all ex- ally deep dive during a long trip. Those data are cept two (one for eachspecies) were recoveredafter comparedwith depths measuredin Cory's Shear- one day. I encounteredan unusual problem in the water (Calonectrisdiomedea) and White-chinned Pe- form of tiny ants that crawled inside the tubes and trel (Procellariaaequinoctialis), which share many removedthe sugar.Ants removedsugar unevenly, in similaritieswith Puffinusbut seemless morphologi- contrastto the sharpboundary created by water,and cally adaptedto diving (Warham1977, 1990). so with careful inspectionI was ableto rejectgauges All five Puffinus shearwaters demonstrated re- affectedby ants. markable diving abilities (Fig. 2). Their maximum Results.--Outof 23 undamagedgauges recovered diving depthsclustered close to the allometricequa- from Wedge-tailedShearwaters, 19 (83%) indicated tion derived from maximum diving depthsof pen- somediving activity (Fig. 1) and their mean maxi- guinsand alcids(maximum depth (meters)= 75.905 mum depthwas 14 m (SD = 23 m, range1-66 m). All M 0'316,where M is body mass in kilograms; Burger 7 gaugesrecovered from Audubon'sShearwaters in- 1991).Maximum depths of Calonectrisand Procellaria dicated diving, and the mean maximum depth was fell well below that regression.The diving abilitiesof 15 m (SD = 12 m, range 6-35 m). Most Wedge-tailed Puffinusshearwaters require more detailed study, us- Shearwaters remained within 20 m and most Au- ing TDR and activitydata-loggers, to confirmwheth- dubon'sShearwaters within 10 m of the surface(Fig. er deep diving forms a significantpart of their for- 1). aging routine. It will be fascinatingto discover Discussion.--Manyprocellariiform species dive to whether they show convergentanatomical, physio- captureprey, but mostspecies remain within 1-5 m logical, and behavioral adaptationsfor prolonged of the surface (Warham 1990, Prince et al. 1994, Hedd diving to thosebeing revealedfor penguins,alcids, et al. 1997),with the exceptionof the specialiseddiv- and cormorants. ing petrels, Pelecanoididae(Prince and Jones1992, Kuroda(1954) regarded smaller species of Puffinus Chastel 1994, Zavalaga and Jahncke1997). Some as morphologicallymore specialized for diving than shearwatersof the genusPuffinus regularly use pur- most other shearwaters. Visual observations of small suit diving for foraging (Brown et al. 1978, 1981; shearwaters (Brown et al. 1978), including Audu- Warham 1990). They show adaptationsfor under- bon'sShearwater (Harris 1969),confirmed that they water swimming, including flattened tarsi and hu- dived, but this study is the first to confirm that div- meri, and shorter wings and higher wing-loading ing occursregularly (in all foragingtrips madein my than most otherpetrels (Kuroda 1954, Warham 1977, small sample), and that they are proficient, deep 1990).Maximum diving depthshave been measured divers. using gaugesin five speciesof Puffinus(Table 1). Keitt et al. (2000) speculated that shearwaters Some published depth records unavoidablycame might be most efficientat diving during wing molt, from prolonged foraging trips. There is a greater when the reduced wing area would produce suffi- chance that those gauges overestimate maximum cient propulsionbut less drag. None of the shear- depthswith repeatedsubmersion and prolongedde- waters in my sample showedany sign of primary July2001] ShortCommunications 757 Penguin-Alcidregresmon Puff'mus••S 0 • •v AU Procellaria ß ß Calonectns 1 1 10 Bodymass (kg) FIG. 2. Puffinusshearwaters had maximum div- ing depths similar to those of alcids and penguins. This log-logplot of maximum diving depthsagainst body masscompares five speciesof Puffinusshear- waters (squares),including Audubon's(AU), Black- vented (BV), Sooty (SO), Short-tailed (ST), and Wedge-tailed(WT) shearwaters,with Cory'sShear- water (triangle) and White-chinned
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