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Foraging Trip Duration Increases for Humboldt Penguins Tagged with Recording De7ices

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Foraging Trip Duration Increases for Humboldt Penguins Tagged with Recording De7ices Griffiths, R., Double, M. C., Orr, K. and Dawson, R. J. G. Piersma, T. 1997. Do global patterns of habitat use and 1998. A DNA test to sex most birds. – Mol. Ecol. 7: migration strategies co-evolve with relative investments in 1071–1075. immunocompetence due to spatial variation in parasite Hamilton, W. D. and Zuk, M. 1982. Heritable true fitness and pressure? – Oikos 80: 623–631. bright birds: a role for parasites? – Science 218: 384–386. Poulin, R. 1996. Sexual inequalities in helminth infections: a Harper, D. G. C. 1999. Feather mites, pectoral muscle condi- cost of being a male? – Am. Nat. 147: 287–295. tion, wing length and plumage coloration of passerines. – Price, P. W. 1980. Evolutionary biology of parasites. – Prince- Anim. Behav. 58: 553–562. ton University Press, Princeton. Hudson, P. J. and Dobson, A. P. 1997. Host-parasite pro- Proctor, H. and Owens, I. 2000. Mites and birds: diversity, cesses and demographic consequences. – In: Clayton, D. parasitism and coevolution. – Trends Ecol. Evol. 15: 358– H. and Moore, J. (eds). Host-parasite evolution: general 364. principles and avian models. Oxford University Press, Ox- Rosen, S., Hadani, A. and Perlstein, Z. 1988. The occurrence ford, pp. 128–154. of Megninia hologastra (Analgidae Gaud, 1974) on poultry in Israel. – Avian Pathol. 17: 921–924. Kirkpatrick, C. E., Robinson, S. K. and Kitron, U. D. 1991. Rozsa, L. 1997. Wing-feather mite (Acari: Proctophyllodidae) Phenotypic correlates of blood parasitism in the common abundance correlates with body mass of passerine hosts: A grackle. – In: Loye, J. E. and Zuk, M. (eds). Bird-parasite comparative study. – Can. J. Zool. 75: 1535–1539. interactions: ecology, evolution and behaviour. Oxford Siikama¨ki, P., Ra¨tti, O., Hovi, M. and Bennett, G. F. 1997. University Press, Oxford, pp. 344–358. Association between haematozoan infections and repro- Komdeur, J. 1991. Cooperative breeding in the Seychelles duction in the pied flycatcher. – Funct. Ecol. 11: 176–183. warbler. – PhD thesis, University of Cambridge, Cam- Thompson, C. W., Hillgarth, N., Leu, M. and McClure, H. E. bridge. 1997. High parasite load in house finches (Carpodacus Komdeur, J. 1992. Importance of habitat saturation and terri- mexicanus) is correlated with reduced expression of a sexu- tory quality for evolution of cooperative breeding in the ally selected trait. – Am. Nat. 149: 270–294. Seychelles warbler. – Nature 358: 493–495. Walter, D. E. and Proctor, H. C. 1999. Mites: ecology, evolu- Larcher, W. 1991. Physiological plant ecology: corrected print- tion and behaviour. – University of New South Wales ing of the second edition. – Springer-Verlag, Berlin. Press, Sydney. May, R. M. and Anderson, R. M. 1978. Regulation and Walther, B. A. and Clayton, D. H. 1997. Dust-ruffling: a stability of host-parasite population interactions. II. Desta- simple method for quantifying ectoparasite loads of live bilizing processes. – J. Anim. Ecol. 47: 249–267. birds. – J. Field Ornithol. 68: 509–518. JOURNAL OF AVIAN BIOLOGY 32: 369–372. Copenhagen 2001 Foraging trip duration increases for Humboldt Penguins tagged with recording de7ices Sabrina S. Taylor (correspondence) and Marty L. Leonard, Department of Biology, Dalhousie Uni7ersity, Halifax, NS, B3H 4J1, Canada. E-mail: [email protected] Daryl J. Boness, National Zoological Park, Smithsonian Institution, Washington, D.C., 20008, USA. Patricia Majluf, Wildlife Conser7ation Society, Armendariz 445, Lima 18, Peru. Marine animals equipped with data recording devices sometimes such as satellite tags, radio-transmitters, and time-depth show changes in their foraging behaviour, which can indicate an recorders are increasingly used to study aspects of adverse effect of the instrument. We attached time-depth recorders (TDRs) and radio-transmitters to study foraging be- foraging behaviour including foraging trip duration, haviour in Humboldt Penguins Spheniscus humboldti and ob- foraging location, and dive characteristics. Despite the served an increase in the duration of foraging trips. We suggest valuable information that these instruments collect, that further tagging of this endangered species be approached their deployment on some species can have negative with caution since this increase in trip duration may negatively affect reproductive and foraging success. effects on foraging behaviour and reproductive success (Wilson et al. 1986, Davis and Miller 1992, Watanuki et al. 1992, Hull 1997). Research on the impact of instruments on behaviour and reproductive success of penguins has produced Foraging behaviour in penguins is difficult to observe equivocal results. Several researchers have found no directly because individuals typically travel large dis- significant effect of instrument deployment on trip du- tances at sea or spend considerable time below the ration (Croxall et al. 1988, Williams et al. 1992, surface (Wilson et al. 1986). Consequently, instruments Watanuki et al. 1992, Wienecke and Robertson 1997) JOURNAL OF AVIAN BIOLOGY 32:4 (2001) 369 whereas others have found a variety of effects including reduce wake and thus drag (Croll et al. 1991). TDRs increased foraging trip duration, decreased foraging were deployed to study foraging behaviour together efficiency, decreased fat levels, or a greater incidence of with radio-transmitters to relocate penguins in the nest abandonment (Gales et al. 1990, Hull 1997, Kirk- event that they deserted their nest. The TDRs were two wood and Robertson 1997, Pu¨tz et al. 1998). The sizes: 6.5×1.8×2.0 cm (37 g) and 8.5×1.1×2.0 cm variable effect of instruments is presumably a function (27.5 g), and the radio-transmitters were 4.0×1.0×2.5 of differences across studies in device size and shape, cm (15 g; dimensions given as length×height×width the method of attachment, and the body condition of for all devices). The sides of the TDRs were tapered to the instrumented animal (Watanuki et al. 1992). It is form a point and the dorsal surface of the radio-trans- therefore difficult to predict the effects of a particular mitters was angled down to form a wedge. Antennas on instrument, especially for species in which instrument the radio-transmitters were 12.5 cm long, 2 mm in deployment is new. However, because instruments po- diameter, and were on angle of 50–60° from horizontal. tentially impact foraging behaviour and reproductive The deployment of TDRs together with radio-transmit- success, it is important to assess their effect on a study ters increased a Humboldt Penguin’s cross-sectional by study basis. This is particularly important for work area by approximately 2.7% and 2.1%. TDR type (high requiring the tagging of endangered species. or low profile) was haphazardly assigned to each pair. We attached time-depth recorders (TDRs) in combi- Penguins were re-captured after approximately two nation with radio-transmitters to endangered Hum- weeks (x¯ 9SD=16.892.11 d, range=10–19 d, n= boldt Penguins Spheniscus humboldti to examine details 23) to minimize the potential effects of chronic expo- of their foraging behaviour (Taylor 2000). In this study sure to drag (Wilson et al. 1989). we compare the length of foraging trips made by tagged The TDRs were programmed to record depth every 7 and untagged penguins to determine the effect of these s while a penguin was in contact with salt-water and to instruments on the birds. record the number of dry readings when the instrument was dry. Using the wet and dry times, we could calcu- late the duration of each foraging trip. Foraging trips Methods were defined as any period at sea which included at least one period of diving activity and ended when the This study was conducted at Punta San Juan, Peru (15° bird returned to land. Humboldt Penguins exhibited 22% S, 75° 12% W), between 21 May and 24 November two types of foraging trip hereafter referred to as day 1999. Punta San Juan is a peninsula safeguarded from trips and overnight trips. If a bird went on a foraging predators by a 1.2 km long concrete wall. Within the trip and returned on the same calendar day, we headland, there are several distinct colonies of breeding classified it as a day trip. If a bird returned on the next Humboldt Penguins. Although most penguins are sur- calendar day, we classified the trip as an overnight trip. face nesters, there are also individual crevice and bur- row nests located around the peninsula and interspersed among and within the colonies (Schwartz et al. 1999). Untagged birds At a study colony of predominantly surface nesting birds, we determined the duration of an untagged indi- Tagged birds vidual’s foraging trip by using arrival and departure Twenty-seven penguins, consisting of 13 pairs and one times from the colony during chick rearing. Individuals individual (n=14 nests), were caught inside individual were recognized by the unique spotting patterns on crevice or burrow nests when their chicks were approx- their breasts (Scholten 1989). To match tagged and imately two to six weeks old and tagged with Mark7 untagged birds, we used observations from untagged time-depth recorders (TDRs; Wildlife Computers, Red- birds (n=30; males=15, females=15) beginning when mond, WA) and VHF radio-transmitters (Advanced their chicks were two weeks old and continuing until Telemetry Systems, Isanti, MN). Radio-transmitters fledging. Observers were present in a blind during all were glued laterally to the TDRs prior to deployment daylight hours (6:00–18:00 hours) and recorded depar- and then both instruments were glued as a unit to the ture and arrival times of untagged birds. To keep the penguin using 5-minute epoxy. The instruments were period of measurement consistent for tagged and un- glued to the dorsal feathers above the uropygial gland tagged birds, we excluded trips (71/157) in which the and below the mid-point of the penguin’s back to departure or arrival times for tagged birds were before minimize drag (Bannasch et al.
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