1002 SHORT COMMUNICATIONS

The Condor 99:1002-1004 0 The Cooper Omlthological Society 1997

MAXIMUM DIVE DEPTHS OF THE PERUVIAN DIVING-PETREL’

CARLOS B. ZAVALAGA Av. Guardia Chalaca 1333, Callao, , e-mail: [email protected]

JAIME JAHNCKE~ Institute de1 Mar de1 Peru ’ (IMARPE), Apartado 22, Callao, Peru, e-mail: [email protected]

Abstract. The mean maximum dive depth attained METHODS for 22 adult Peruvian Diving-petrels (Pelecanoides The field work was carried out on La Vieja Island, garnotii) was 31.6 + 3.6 m. The deepestdive was 83.1 Peru (14”16’S, 76”ll’W) from August 1995 to May m. This maximum dive depth was 81% deeper than 1996. The island lies at the entranceof Independencia that predicted by its body size (46 m) and the deepest Bay, about 50 km south of the ParacasPeninsula. reported for a weighing less than 210 g. Depth gauges were deployed on 66 adult at Key words: Peruvian Diving-petrels, diving different stagesof the breeding cycle (incubation and depths, foraging behavior, Pelecanoidesgamotii. chick rearing). Adults were capturedin their burrows, weighed, measured,and ringed. Gaugeswere attached to feathersin the center of the ’s back using quick- The maximum dive depth of several taxa of free-living set epoxy glue (Devcon 5-minute epoxy). The devices seabirdshas been successfullydetermined using cap- were retrieved after l-3 days. illary-tube gauges. Although these devices provide Capillary-tube depth gauges, similar to those de- only the record of the deepest dives, which normally scribed by Burger and Wilson (1988), consistedof a 6 represent less frequent exploratory dives beyond the cm length of flexible, clear PVC tubing (internal di- ’s normal foraging depths (Lishman and Croxall ameter of 0.8 mm), coated internally with a thin layer 1983. Piatt and Nettleshio 1985. Watanuki et al. 1996). of indicator powder (one part methylene blue:three they are useful in that they provide an initial insight parts talcum powder). Once coated, the tube was heat- into an animal’s underwater capabilities, habitat use, sealed at one end. Each tube weighed approximately and accessto deep prey (Adams and Brown 1983, Hax- 0.5 g, which was < 0.25% of adult body weight (X = ris et al. 1990. Prince and Jones 1992). Maximum 202 g, n = 65), and had a cross-sectionalarea of less depth gaugesare light-weight and representat present than 10 mm2, about 0.7% of the maximum cross-sec- the only viable and reliable technique to measure the tional area of an adult bird (X = 14.6 cm2, n = 32). diving capabilities of small birds such as diving-pet- During diving, water enters the tube by hydrostatic rels, which are incapable of carrying heavy and more pressure,compressing the volume of air inside the lu- sophisticateddevices. men. The volume of compressedair varies inversely The Peruvian Diving-petrel (Pelecunoides garnotii), the largest of the four extant diving-petrels,is endemic to the cold nutrient-rich Peruvian Coastal Current (Murphy 1936). Once abundant,these birds have been adversely affected by hunting, fishing activities, and reduction of nesting habitat (Murphy 1936, Hays 1989). Although Peruvian Diving-petrels are consid- ered endangered(Wege and Long 1995), information about the breeding and foraging ecology of thesebirds is scarce. The maximum breeding population in Peru was estimated to be 13,000 pairs in 1996, whereas their distribution was restricted to only two islands in central Peru: La Vieja and San Gallan (Jahncke and Goya, unpubl. data). In this study we report maximum dive depths at- tained by breeding Peruvian Diving-petrels and com- *L pare them with data obtained from other wing-pro- 0 pelled divers. Maximum Depth (m) FIGURE 1. Maximum dive depths of Peruvian Div- ’ Received 15 April 1997. Accepted 30 July 1997. ing-petrels (n = 22) breeding in La Vieja Island, Peru, * Current address:Apartado 18-0807, Lima 18, Peru. between August 1994 and May 1995. SHORT COMMUNICATIONS 1003

TABLE 1. Maximum diving depths recorded for three species of diving-petrels and two similar-sized alcids. D,, predicted was determined by the equation D,, = 75.905M”-3’6 (Burger 1991).

Weight MClll Observed Predicted Species (9) D,, (ml %, Cm) D,, Cm) n Reference Diving-petrels Pelecanoides georgicus 93.0 25.7 48.6 35.8 6 Prince and Jones 1992 P. urinatrix 147.4 38.9 63.6 41.1 103 Chaste11994 P. garnotii 202.0 31.6 83.1 45.6 22 This study Alcids Ptychoramphus aleuticus 190.0 - 43.0 44.9 23 Burger 1991 Alle alle 160.0 - 35.0 42.5 - Burger 1991

with hydrostatic pressure (with increasing depth) as depths of penguins and alcids were positively corre- described by Boyle’s Law. When the bird is at the lated to body size and can be predictedby the equation surface, the water washes out the powder, leaving a D mar= 75.905M03r6,where M = mass in kg, for both line at the point of maximum compression.The max- groups combined. Using this equation with the mean imum depth attained was calculatedby the equation: weight of Peruvian Diving-petrels (202 g), a maximum dive of 45.8 m is expected. This value is about half D max= lO.O8[(L&) - 11; that of the observed maximum dive of 83.1 m. Like- where D,, is the maximum depth in meters, 10.08 is wise, maximum depths attainedby the three speciesof the height (m) of a column of sea water equivalent to diving-petrels were deeper than those recorded for 1 atmosphereof pressure,L, is the total length of the similar-sized alcids (Table l), which are recognized to tube, and Ld is the length of the tube with undissolved be close ecological and morphological analogues of indicator (Burger and Wilson 1988). After recovery of diving-petrels (Thoresen 1969). This evidence suggests the recorders,the length of the tube with indicator was that Peruvian Diving-petrels are capable of exploring measuredto the nearest 0.1 mm using calipers. Tubes deep waters and that diving capabilities of diving-pet- with water droplets in the lumen were discarded. rels may differ from those exhibited by other wing- The accuracy of the depth gauges was checked by propelled divers. lowering nine of them into sea-water from the SNP-1 Diving patterns of Peruvian Diving-petrels may be Research Vessel of IMARPE (Instituto de1 Mar de1 variable-according to location, time of the year,-and Peru) to known depths ranging from 10 to 150 m. behavior of its prey (Wilson and Wilson 1990). The Means are expressed5 SE. major prey speciesconsumed by thesebirds aroundLa Vieja Islandduring the study period were small crus- RESULTS taceans,mainly euphausiids(25.2% by mass) and lar- Maximum depth gaugesoverestimated real depths by val stages of Peruvian anchovies (Engrazdis ringens) an average of 13.2% (2.2-35.0%) with shallower and other (27.3% by mass) (Jahnckeet al., un- depths (< 70 m) being recordedmore precisely (< 8% publ. data). Euphausiidsare widespread to depths of error) than deeper depths (> 70 m). Estimated depths at least 300 m (Brinton 1962, cited in Santanderet al. could be correctedby the equation:Real depth = 3.12 1981), whereas Peruvian anchovy larvae are confined + 0.8 X estimateddepth, for estimateddepths ranging to the upper 60 m (Sameoto 1981). Both prey types from 10 to 150 m (+ = 0.99, n = 9). show die1 vertical migration, remaining in deeper wa- Forty-one gauges (62%) were recovered after one ter strata during the day and rising to the surface by foraging trip, but only 22 gave readablemeasurements. night (Brinton 1962, cited in Santander et al. 1981, Mean maximum dive depth was 31.6 ? 3.6 m, and Sameoto 1981). Because Peruvian Diving-petrels are 91% of the maximum dives were between 10 and 50 mainly diurnal foragers (Jahncke, unpubl. data), it is m (Fig. 1). The shallowest dive was 10.0 m, and the expectedthat these birds need to reach deep water stra- deepestdive was 83.1 m. There was no significantdif- ta to catch food. ference in the maximum dive depth attained by birds We thank Elisa Goya, Domenica Zileri, Raul San- incubating eggs (n = 34.4 C 3.8 m, n = 5) and those chez, Cecilia Flores, and Margaret Fraserfor their help rearing chicks (x = 30.8 I 4.6 m, n = 17) (Mann- with the field work. David Duffy, Rory Wilson, Ian Whitney U = 28, P = 0.27). Stirling, Patricia Majluf, and two anonymousreview- DISCUSSION ers suggested improvements to early drafts of this manuscript.We are grateful to PESCA PERU for per- The mean maximum dive depth attained by the Peru- mission to work on the island and for providing ac- vian Diving-petrel was similar to those reported for commodations. other species of diving-petrels (Table 1). However, when the deepestdive depthsrecorded for each species LITERATURE CITED are compared, values are related, among other things, to body size, as in other species of diving ADAMS, N. J., AND C. R. BROWN. 1983. Diving depths (Piatt and Nettleship 1985, Burger 1991). of the Gentoo Penguin (Pygoscelis Papua). Con- Burger (1991) determined that maximum dive dor 85:503-504. 1004 SHORT COMMUNICATIONS

BURGER,A. E. 1991. Maximum diving depths and un- Zecanoides georgicus at Bird Island, South Geor- derwater foraging in alcids and penguins. Can. gia. Antarct. Sci. 4:433-434. Wildl. Serv., Occas. Pap. 68:9-15. SAMEOTO,D. 1981. Distribution and abundanceof six BURGER,A. E., AND R. P WUON. 1988. Capillary- speciesof larvae in Peruvian waters and their tube depth gauges for diving : an assess- relationship with the physical and biological en- ment of their accuracy and applicability. J. Field vironment: Boletin Inst&uto de1 Mar de1 Peru. Omithol. 59:345-354. Volumen Extraordinario:164-170. CHASTEL,0. 1994. Maximum diving depths of Com- SANTANDER,H., G. LUYO,S. CARRASCO,M. V~LU, AND mon Diving Petrels Pelecanoides urinatrix at Ker- 0. S. DE CASTILLO.1981. Catalog0 de zooplanc- guelen Islands. Polar Biol. 14:211-213. ton en el mar peruano.Primera parte: area Pisco- HARRIS,M. P, H. TOWLL,A. E RUSSELL,AND S. WAN- San Juan. Boletfn Instituto de1Mar de1Peru 6:1- LESS. 1990. Maximum dive depths attained by 75. THORESEN,A. C. 1969. Observationson the breeding auks feeding young on the Isle of May, Scotland. behaviourof the Pelecanoides u. ur- Scottish Birds 16:25-28. inatrix (Gmelin). Notomis 16:241-260. HAYS, C. 1989. The Peruvian Diving-petrel in Peru. WATANUKI,Y., A. KATO,AND Y. NAI~O. 1996. Diving Oryx 23:102-105. performance of male and female JapaneseCor- LISHMAN,G. S., AND J. F! CROXALL. 1983. Diving morants. Can. J. Zool. 74: 1098-l 109: depths of the Chinstrap Penguin, Pygoscelis ant- WECE D. C.. AND A. J. LONG. 1995. Kev areas for arcticu. Br. Antarct. Surv. Bull. 61:21-25. threatenedbirds in the Neotropics. BGdlife Con- MURPHY,R. C. 1936. Oceanic birds of South Amer- servation Series No. 5. Smithson. Inst. Press, ica. Vol. 2. Macmillan, New York. Washington,DC. PIAT~,J. E, AND N. NETIZESHIP.1985. Diving depths WILSON,R. l?, AND M. T. Wnso~. 1990. Foraging of four alcids. Auk 102:293-297. ecology of breeding Spheniscus penguins,p. 18l- PRINCE, P A., AND M. JONES.1992. Maximum dive 206. In L. S. Davis and J. T Darby [eds.], Penguin depthsattained by South Georgia diving petrel Pe- biology. Academic Press, San Diego.

The Condor 99:lOCb1007 0 The CooperOrnithological Society 1997

DIVING DEPTHS OF TWO TROPICAL PELECANIFORMES: THE RED-TAILED TROPICBTRD AND THE RED-FOOTED BOOBY’

MAITHIEU LE CORRE CEBC-CNRS Villiers en Bois-79360 Beauvoir sur Niort, France

Abstract. The diving ability of two tropical Pele- Most boobies (tropical Szdidae) and all tropicbirds caniformes, the Red-footed Booby (Sulu sulu) and the (Phaethontidue) a& pelagic plunge diving seabirds Red-tailed Tropicbird (Phaethon rubricauda) was (Ashmole 1971, Nelson 1978, Schreiber and Clapp studiedon EuropaIsland, southernMozambique Chan- 1987), foraging solitarily (tropicbirds,Masked Booby) nel, using capillary depth recorders fitted on breeding or collectively-(most boobies), and feeding mainly on adults. Both speciesmainly exploited the first 4 m of flying fishes and squid (Ashmole and Ashmole 1967, the water column. Although such a depth can be Diamond 1974, 1975, Harrison et al. 1983). Plunge reached solely by passive plunge diving, the range of divers supposedlyhave limited diving capacities, the depths reached by the two species suggeststhat they depth they reach depending mainly on the momentum may, at least sometimes, use their feet and wings to gained during the plunge (Ashmole 1971). However, perform active underwater pursuit swimming. Intra- at least Cape Gannets (S&I cupensis) can use their specific comparisonof the depth reachedby Red-tailed wings and feet to gain additional depth (Adams and Tropicbirdsalso suggeststhat this speciesmay change Walter 1993). Here I report diving performances of its diving behavior seasonally. two tropical plunge divers, the Red-tailed Tropicbird (Phaethon rubricaudu, 0.8 kg) and the Red-footed Key words: diving depth, feeding ecology, Red- Booby (Sula sula, 0.9 kg). tailed Tropicbird, Red-footed Booby, Phaethon rubri- cauda, Sula sula, Europa Island. METHODS The study was conductedon Europa Island (22”2O’S, 40”22’E) in the southernMozambique Channel. About 1Received 8 May 1997. Accepted 31 July 1997. 3,000 pairs of Red-footed Boobies and about 3,500