263 Site and Mate Choice in Seabirds: An Evolutionary Approach Joël Bried and Pierre Jouventin CONTENTS 9.1 Introduction 263 9.2 The Major Evolutionary Constraint in Seabirds: Breeding on Land and Feeding at Sea 269 9.2.1 The Key Factor 269 9.2.2 Phylogenetic Constraints 272 9.2.3 Phenological Constraints 272 9.3 Habitat Selection 278 9.3.1 Choice of the Breeding Place 278 9.3.2 Nest-Site Selection 279 9.4 Mate Choice 279 9.5 Site and/or Mate Tenacity, or Switching? 285 9.5.1 Benefits of Site and Mate Fidelity 285 9.5.2 Costs of Site and Mate Fidelity 287 9.5.3 Benefits and Costs of Divorce and Site Changes 288 9.5.4 Changing Site and/or Mate 288 9.6 Conclusions and Perspectives 291 9.6.1 Which Strategy Seems the Most Adaptive for Seabirds? 291 9.6.2 Is Fidelity Positively Related to Longevity? 291 9.6.3 Is Mate Fidelity Just a By-Product of Nest Fidelity? 293 9.6.4 Influence of Breeding Success on Fidelity: Relevance to Conservation 294 Acknowledgments 295 Literature Cited 295 9.1 INTRODUCTION In more than 90% of avian species, monogamy is the mating system (Lack 1968) but it still remains “the neglected mating system” (Mock 1985), because many studies on mating systems focus on the evolution and the maintenance of alternative mating systems (i.e., polygamy and promiscuity) rather than on the reasons why monogamy evolved. The concept of monogamy has been debated by (among others) Wickler and Seibt (1983) and Gowaty (1996). Distinguished from genetic monogamy, social monogamy can be defined as the association of one male and one female usually with some level of biparental care. In birds, this partnership, exclusive for incubation and chick- rearing, can be maintained during an entire lifetime. 9 © 2002 by CRC Press LLC 264 Biology of Marine Birds The choice of a breeding place (Cody 1985, Ens et al. 1995) and a sexual partner (Orians 1969, Hunt 1980, Ligon 1999) has important consequences for reproduction. In birds, males classically compete over sites and/or females, whereas females perform mate choice (Darwin 1871, Orians 1969, Trivers 1972). Whether individuals should retain their site and/or mate from year to year, or change, is ultimately determined by breeding success, considering both previous and expected future reproductive performances (Greenwood and Harvey 1982, Switzer 1993, McNamara and Forslund 1996). According to Hinde (1956) and Rowley (1983), individuals of long-lived species should be able to retain both site and mate from year to year, because of their high adult survival rates. Moreover, life history theory predicts that high longevity should be associated with reduced fecundity or low reproductive effort (Stearns 1992). Therefore, individuals of long-lived species should optimize their reproductive outputs, while minimizing the costs of breeding not to jeopardize their future survival and residual reproductive value (Drent and Daan 1980, Partridge 1989, Ricklefs 1990, Stearns 1992; but see also Erikstad et al. 1998). Maximizing their chances to replace themselves (by producing at least one chick that will recruit into the breeding population) can be achieved through a high number of breeding attempts (iteroparity), and hence a long reproductive life span. Because site and mate fidelity are known to enhance reproductive performances in many avian species (Domjan 1992, Ens et al. 1996), long-lived species classically are expected to show high site and mate fidelity, with fidelity rates and life expectancy being positively correlated (Rowley 1983; Figure 9.1). However, very few studies have so far examined the relationships between fidelity and survival (e.g., Ens et al. 1996, Bried, Pontier, and Jouventin in preparation), considering fidelity rates as demographic parameters. Seabirds appear as a choice model for these studies, being particularly long-lived, laying small clutches, and having a deferred sexual maturity (Jouventin and Mougin 1981; see also Chapter 5 by H. Weimerskirch; Table 9.1). Furthermore, the probability for seabird young to recruit into the breeding population is low (review in Nelson 1980; see also Ollason and Dunnet 1988, Wooller et al. 1989, Weimerskirch et al. 1992, Prince et al. 1994, Weimerskirch and Jouventin 1997), because a high proportion of seabird fledglings die from starvation during their first year of independence, presumably lacking sufficient foraging skills (Nelson 1980, Nur 1984). Due to biparental care, all seabird species are socially monogamous (Lack 1968). However, genetic monogamy may not always occur, promiscuous matings and polygyny having been observed in FIGURE 9.1 A Short-tailed Albatross incubating its egg on Torishima Island, Japan. The incubation period is about 60 days and it takes the pair about 180 days to raise their single chick. (Photo by E. A. and R. W. Schreiber.) © 2002 by CRC Press LLC Site and Mate Choice in Seabirds: An Evolutionary Approach 265 TABLE 9.1 Nest and Mate Fidelity in Seabirds Taxon a Locality Nest Fidelity b (%) Mate Fidelity b (%) S (ALE) Body Mass (g) References c Sphenisciformes Aptenodytes patagonicus Iles Crozet 39.4 (site fidelity) 22.4 0.952 (21.33) 13,400 1, 2, 3, 4 A. patagonicus South Georgia — 18.8 — 13,800 5 A. forsteri Terre Adélie — 14.5 0.91 (11.61) 30,000 2, 6, 4 Pygoscelis adeliae Cape Crozier 59.4 18–50 0.696 (3.79) 3,900 7, 7, 8, 4 P.adeliae Cape Bird 98.2 56.5 0.736 (4.29) 4,200 9, 9, 9, 4 P. adeliae Wilkes Land 76.8 84.0 0.77 (4.85) 4,490 10, 10, 10, 4 P. papua papua Iles Crozet — 76.0 0.865 (7.91) 6,740 11, 8, 4 P. p. papua South Georgia 93.0 90.2 ca. 0.8 (ca. 5.5) 6,800 12, 12, 13, 4 P. p. ellsworthi King George Is. 61.4 90.0 — 5,300 14, 14, 14, 15 P. antarctica King George Is. 87.9 82.0 — 4,000 14, 14, 14, 4 Eudyptes (chrysolophus) chrysolophus South Georgia 83 90.8 — 4,120 12, 12, 16 E. (c.) schlegeli Macquarie Is. — 80.0 0.86 (7.64) 5,000 16, 17, 18 E. chrysocome filholi Iles Kerguelen 53.0 78.6 — 2,500 19, 19, 19 E. c. moseleyi Amsterdam Is. 34.9 46.3 0.84 (6.75) 2,400 19, 19, 20, 21 E. robustus The Snares — no case reported — 3,100 4, 4 Megadyptes antipodes New Zealand 30.0 82.0 0.87/0.86 (ca. 8.5) 5,300 22, 23, 24, 4 Spheniscus mendiculus Galápagos Is. — >89 0.844 (6.91) 2,030 25, 16, 16 S. demersus South Africa 59.8 86.2 0.617 (3.11) 3,100 26, 26, 26, 16 S. magellanicus Punta Tombo 80/70 90.4 0.85 (7.17) 4,440 27, 16, 16, 16 Eudyptula minor Philip Is. 43.9 82.0 0.858 (7.54) 1,110 28, 28, 28, 4 Procellariiformes Diomedea exulans South Georgia 20.0 no case reported 0.94 (17.17) 8,700 29, 30, 4 D. exulans Iles Crozet 28.9 95.1 0.931 (14.99) 9,600 31, 19, 32, 33 D. amsterdamensis Amsterdam Is. — 97.9 0.966 (29.91) 6,270 19, 19, 34 D. epomophora epomophora Campbell Is. — no case reported — 9,280 35, 4 D. e. sanfordi Taiaroa Is. — 1 case reported 0.946 (19.02) 6,500 36, 36, 4 Diomedea (Phoebastria) irrorata Galápagos Is. — no case reported 0.95 (20.50) 3,500 37, 37, 37 D. immutabilis Midway Atoll — 97.9 0.947/0.946 (ca. 19.19) 2,900 38, 39, 40 © 2002 by CRC Press LLC 266 Biology of Marine Birds TABLE 9.1 (Continued) Nest and Mate Fidelity in Seabirds Taxon a Locality Nest Fidelity b (%) Mate Fidelity b (%) S (ALE) Body Mass (g) References c Phoebetria fusca Iles Crozet 41.1 94.8 0.95 (20.50) 2,600 19, 19, 41, 42 Diomedea (Thalassarche) chlororhynchos Amsterdam Is. 92.6 90.6 0.912 (11.86) 2,100 43, 19, 41, 42 D. bulleri The Snares 67.0 96.2 0.913 (11.99) 2,700 44, 44, 45, 4 D. chrysostoma Campbell Is. — 96.3 0.953 (21.78) 3,180 46, 47, 4 D. melanophris melanophris Iles Kerguelen 74.1 92.3 0.914 (12.13) 3,740 19, 19, 19, 4 D. m. melanophris South Georgia 93.5 — 0.934 (15.65) 3,600 48, 48, 4 D. m. impavida Campbell Is. — 95.5 0.945 (18.68) 2,900 46, 47, 4 Pagodroma nivea Terre Adélie 89.8 88.3 0.934 (15.65) 380 49, 49, 50, 51 Daption capense capense Terre Adélie 88.0 85.0 — 472 52, 52, 53 D. c. capense South Orkney Is. 84.0 73.0 0.942 (17.74) 425 54, 54, 54, 55 D. c. australe The Snares 97.5 97.3 0.892 (9.76) 435 56, 56, 56, 57 Fulmarus glacialoides Terre Adélie 82.5 77.1 0.916 (12.40) 800 19, 19, 19, 4 F. glacialis Orkney Is. 93.4 96.9 0.968 (31.75) 813 58, 58, 59, 60 Macronectes giganteus Terre Adélie 59.0 80.8 0.902 (10.70) 4,500 19, 19, 51, 51 M. giganteus South Orkney Is. 92.9 no case reported — 4,360 61, 61, 61 Pelecanoides urinator Iles Kerguelen 81.6 92.8 0.807 (5.68) 140 19, 19, 19, 19 Pterodroma lessonii Iles Kerguelen 96.6 91.2 0.921 (13.16) 708 19, 19, 51, 62 P. macroptera Iles Kerguelen 80.2 87.5 — 581 19, 19, 63 P. inexpectata The Snares 96.9 >83 — 320 64, 64, 64 P. phaeopygia Galápagos Is. 96.7 — — 410 65, 65 Calonectris diomedea borealis Salvages Is. 91.4 94.0 0.956 (23.23) 890 66, 66, 67, 4 C. d. diomedea Crete 95.9 96.4 0.89 (9.59) 552 68, 68, 68, 69 Puffinus puffinus Skokholm Is. 93.3 90.3 0.905 (11.03) 450 70, 70, 70, 71 P. tenuirostris Bass Strait — 82.2 0.897/0.899 (ca. 10.30) 590 72, 73, 4 Procellaria aequinoctialis Iles Crozet 80.5 93.7 — 1,300 74, 74, 75 P. parkinsoni New Zealand — 88.0 0.94 (17.17) 700 76, 76, 71 P. cinerea Iles Kerguelen 90.2 95.9 0.924 (13.66) 1,131 19, 19, 51, 77 Bulweria bulwerii Salvages Is. 63.0 78.5 0.947 (19.37) 95 78, 78, 79, 80 Halobaena caerulea Iles Kerguelen 88.3 80.0 0.88 (8.83) 190 19, 19, 51, 81 Pachyptila belcheri Iles Kerguelen 87.5 79.2 0.852 (7.26) 145 32, 32, 51, 63 © 2002 by CRC Press LLC Site and Mate Choice in Seabirds: An Evolutionary Approach 267 P. desolata Iles Kerguelen 86.5 88.0 — 150 19, 19, 19 P. turtur Whero Is. 87.0 — 0.844 6.91) 130 80, 82, 4 Oceanites oceanicus South Orkney Is. — 80.0 0.908 (11.37) 40 83, 83, 83, 83 Hydrobates pelagicus Skokholm Is. — 77.3 0.88 (8.83) 28 84, 71, 71 Oceanodroma leucorhoa Maine, U.S.A. 95.0 — 0.86 (7.64) 45 85, 86, 71 Pelecaniformes Phaethon rubricauda Kure Atoll 25.0 — — 612 87, 88 P. lepturus Seychelles Is. — 97.0 — 341 89, 89 Morus bassanus Bass Rock 89.8 83.5 0.89 (9.59) 3,000 90, 90 S. dactylatra personata Kure Atoll 10.0 54.8 0.895 (10.02) 2,030 90, 90, 90, 90 Sula (d.) granti Galápagos Is. 87.1 — 0.832 (6.45) 1,750 90, 87 S. leucogaster Kure Atoll — ≥97.7 0.92—0.955 (ca. 16.50) 1,110 90, 90, 90, 90 S. abbotti Christmas Is. — ≥90 — 1,480 90, 4 Phalacrocorax aristotelis Isle of May 49.2 69.0 0.84 (6.75) 2,000 91, 91, 92, 71 P. penicillatus Farallon Is. 62.3 — 0.80 (5.50) 2,450 93, 93, 71 P. atriceps South Orkney Is. — 40.3 — 2,880 94, 71 Nannopterum harrisi Galápagos Is. 35.9 11.9 0.876 (8.56) 3,200 95, 95, 95, 71 Charadriiformes Catharacta skua skua Foula Is. — 93.6 0.93 (14.78) 1,418 96, 97, 97 C. s. lönnbergi Iles Kerguelen 98.3 96.5 0.925 (13.83) 1,835 19, 19, 19, 19 C. s. lönnbergi Anvers Is. — > 89 0.95 (20.50) 1,700 98, 98, 99 C. maccormicki Terre Adélie 89.0 90.9 0.912 (11.86) 1,405 19, 19, 19, 100 C. maccormicki Anvers Is. — > 85 0.95 (20.50) 1,200 98, 98, 99 C. maccormicki Cape Crozier 87.3 98.5 0.938 (16.63) 1,300 101, 102, 99 L. (novaehollandiae) scopulinus New Zealand — 89.5 0.856 (7.44) 280 103, 103, 99 Rissa tridactyla tridactyla Great Britain — 71.9 0.81/0.86 (ca. 6.56) 408 104, 105, 106 R. t. pollicaris Alaska — 80.7 0.93 (14.78) 408 107, 107, 106 Sterna anaethetus Western Australia 82.3 — 0.78 (5.04) 127 108, 108, 99 Sterna hirundo Germany — 81.1 0.89 (9.59) 134 109, 110, 109 Anous minutus Ascension Is. 81.8 — — 118 111, 111 Uria lomvia Prince Leopold Is. 73.0 — 0.91 (11.61) 945 112, 113, 113 U. aalge Isle of May 85.7 88.3 0.949 (20.11) 862 114, 115, 116, 117 Alca torda Isle of May 93.0 94.3 0.888 (9.43) 710 118, 118, 118, 113 Ptychoramphus aleuticus Farallon Is. — 92.7 0.75 (4.50) 170 119, 119, 113 Cepphus grylle Iceland 90.0 95.5 0.87 (8.19) 500 120, 120, 120, 113 Aethia cristatella Buldir Is. 75/62 64.5 0.89 (9.59) 260 121, 121, 121, 113 © 2002 by CRC Press LLC 268 Biology of Marine Birds TABLE 9.1 Nest and Mate Fidelity in Seabirds Taxon a Locality Nest Fidelity b (%) Mate Fidelity b (%) S (ALE) Body Mass (g) References c A. pusilla Pribilof Is. — 63.6 0.808 (5.71) 85 122, 122, 113 Fratercula arctica Skomer Is. 92.2 92.2 0.942 (17.74) 460 123, 123 123, 113 F. arctica Unknown locality — 84.0 0.87 (8.19) 460 124, 124, 113 a Only adult individuals (i.e., known to have bred in the past) were considered. Data from populations kno wn to live in unstable environments, or not to be in equilibrium, were excluded. b Studies involving less than 25 individual-years or 25 pair-years for site fi delity and mate fidelity, respectively, were excluded. Only adult indi viduals were considered. Site fidelity rates were calculated as 1 minus (number of site changes/number of adult-years). Mate fi delity was calculated as 1 minus the probability of divorce when both previous partners survive, following Black (1996). When two values separated by a slash (/) are gi ven for the same parameter (e.g., 80/70), the former is for males, the latter for females. c Numbers refer to the source of nest fidelity, mate fidelity, adult survival rate, respectively, and when data were available. Although some of these sources did not express fidelity rates in the same manner as ours, they provided the data that enabled us to calculate them as described abo ve. 1, Barrat (1976); 2, Bried et al. (1999); 3, Weimerskirch et al. (1992); 4, Marchant and Higgins (1990); 5, Olsson (1998); 6, J ouventin and Weimerskirch (1991); 7, Ainley et al. (1983); 8, Ainley and DeMaster (1980); 9, Davis (1988); 10, Penne y (1968); 11, Bost and Jouventin (1991); 12, Williams and Rodw ell (1992); 13, Croxall and Rothery (1994); 14, Trivelpiece and Trivelpiece (1990); 15, Volkman et al. (1980); 16, Williams (1995); 17, Carrick (1972); 18, Carrick and Ingham (1970); 19, this study; 20, Guinard et al. (1998); 21, E. Guinard (unpublished data); 22, Richdale (1949); 23, Richdale (1947); 24, Jouv entin and Mougin (1981); 25, Boersma (1976); 26, LaCock et al. (1987); 27, Scolaro (1990); 28, Reilly and Cullen (1981); 29, Tickell (1968); 30, Croxall et al. (1990); 31, Fressanges du Bost and Ségonzac (1976); 32, Weimerskirch and Jouventin (1997); 33, Rice and Kenyon (1992); 34, Jouventin et al. (1989); 35, Waugh et al. (1997); 36, Robertson (1993); 37, Harris (1973); 38, Rice and K enyon (1962); 39, Fisher (1975); 40, Frings and Frings (1961); 41, Weimerskirch et al. (1987); 42, Weimerskirch and Jouventin (1897); 43, Jouventin et al. (1983); 44, Sagar and Warham (1997); 45, P. M. Sagar, J. Molloy, H. Weiberskirch, and J. Warham (unpublished data); 46, S. M. Waugh and J. Bried (unpublished data); 47, Waugh et al. (1999); 48, Prince et al. (1994); 49, Jouventin and Bried (in press); 50, Chastel et al. (1993); 51, Chastel (1995); 52, Mougin (1975); 53, Isenmann (1970); 54, Hudson (1966); 55, Pinder (1 966); 56, Sagar et al. (1996); 57, Sagar (1986); 58, Ollason and Dunnet (1978); 59, Dunnet and Ollason (1978); 60, Ollason and Dunnet (1988); 61, Conro y (1972); 62, Zotier (1990b); 63, Weimerskirch et al. (1989); 64, Warham et al. (1977); 65, Cruz and Cruz (1990); 66, Mougin et al. (1987a); 67, Mougin et al. (1987b); 68, Sw atschek et al. (1994); 69, Ristow and Wink (1980); 70, Brooke (1990); 71, del Hoyo et al. (1992); 72, Bradley et al. (1990); 73, Wooller and Bradle y (1996); 74, Bried and Jouventin (1999); 75, A. Catard (unpublished data); 76, Imber (1987); 77, Zotier (1990a); 78, Mougin (1989); 79, Mougin (1990); 80, Warham (1990); 81, Chastel et al. (1995a); 82, Richdale (1963); 83, Beck and Brown (1972); 84, Scott (1970); 85, Morse and Buchheister (1979); 86, Warham (1996); 87, Harris (1979b); 88, Fleet (1974); 89, Phillips (1987); 90, Nelson (1978); 91, Aebischer et al. (1995); 92, Potts (1969); 93, Boekelheide and Ainley (1989); 94, Shaw (1986); 95, Harris (1979a); 96, Catry et al. (1997); 97, Furness (1987); 98, Pietz and Parmelee (1994); 99, Higgins and Davis (1996); 100, Jouventin and Guillotin (1979); 101, Ainley et al. (1990); 102, Wood (1971); 103, Mills et al. (1996); 104, Coulson (1966); 105, Coulson and Wooller (1976); 106, Burger and Gochfeld (1996); 107, Hatch et al. (1993); 108, Dunlop and Jenkins (1992); 109, González-Solís et al. (1999); 110, Wendeln and Becker (1998); 111, Ashmole (1962); 112, Gaston and Nettleship (1981); 113, Nettleship (1996); 114, Harris et al. (1996); 115, Ens et al. (1996); 116, Harris and Wanless (1995); 117, Cramp (1985); 118, Harris and Wanless (1989); 119, Sydeman et al. (1996); 120, Petersen (1981); 121, Gaston and Jones (1998); 122, Jones and Montgomerie (199 1); 123, Ashcroft (1979); 124, Davidson (unpublished data, in Ens et al. (1996). © 2002 by CRC Press LLC Site and Mate Choice in Seabirds: An Evolutionary Approach 269 gulls (Burger and Gochfeld 1996). Because living organisms tend to optimize their own fitness, but also that of their offspring (Maynard-Smith 1978), do the long-lived seabirds choose their breeding places and their partners carefully? In this chapter, we provide a new insight into the relationships between site fidelity, mate fidelity, and longevity, by using an evolutionary approach with seabirds as a model. In order to achieve this purpose, we will (1) identify the constraints on reproduction faced by seabirds, and (2) test the classical predictions concerning site, mate choice, and fidelity (see above) by assessing the effects of these selective pressures on the reproductive strategy of seabirds. 9.2 THE MAJOR EVOLUTIONARY CONSTRAINT IN SEABIRDS: BREEDING ON LAND AND FEEDING AT SEA 9.2.1 T HE KEY FACTOR Seabirds face an important constraint during reproduction, which appears as the key factor in the evolution of their life history traits: they exclusively rely on marine resources for feeding and yet they need to come ashore for breeding (Jouventin and Mougin 1981). For “inshore” and “offshore” feeders, nesting and feeding areas are not only distinct, but there is a continuous gradient from the more coastal seabirds to the most pelagic ones: foraging trips can range from a few hundred meters from the nest in terns to several thousands kilometers in albatrosses and petrels (Jouventin and Weimerskirch 1991, Weimerskirch 1997, Weimerskirch et al. 1999). Consequently, trips can last up to several days and sometimes several weeks, and their duration has affected the evolution of seabird life histories (see Figures 9.2 and 9.3). These foraging trips represent a constraint in seabirds, for demography but also for morphology (wing shape) and metabolism, because seabirds must fly long distances and fast when on land (Warham 1975, 1990, Schreiber and Schreiber 1993, Chaurand and Weimerskirch 1994a). A breeding adult that undertakes a long foraging trip while its partner incubates or broods will have to return to its nest before its mate has exhausted its body reserves and abandoned the nest, implying a good synchronization between mates (e.g., Jouventin et al. 1983 for albatrosses, Davis 1988 for Adélie Penguins [Pygoscelis adeliae], Schreiber and Schreiber 1993 for Red-tailed Tropicbirds [Phaethon rubricauda]). Nevertheless, successful breeding in seabirds does not only depend on food and synchronization between parents; two other conditions must be met on land. The first one is the ownership of a breeding territory to have a place to incubate the clutch (Newton 1992). The second condition is obtaining the sexual partner. In many seabird species, males return ashore earlier than females at the onset of breeding and settle on their nests before attracting a mate (Hunt 1980; for examples of taxonomic groups, see e.g., Warham 1990 for albatrosses and petrels, and Nelson 1983 for sulids and some cormorants). However, in Emperor Penguins (Aptenodytes forsteri), females generally return earlier than males (Bried et al. 1999), and both sexes can return simultaneously in frigatebirds (Nelson 1983), but also in some terns and alcids (Hunt 1980). However, site quality and mate quality vary in birds, including seabirds (e.g., Adélie Penguins, Carrick and Ingham 1967; Caspian Terns [Hydroprogne caspia], Cuthbert 1985; sulids, Nelson 1988; Snow Petrels [Pagodroma nivea], Chastel et al. 1993). Therefore, individuals should settle on the most suitable sites available (this implies proximity to foraging area, concealment from weather and potential predators, and easy access and departure for breeders) and should obtain as high quality mates as possible (“ideal” choice, Fretwell and Lucas 1970). Because of the constraints of oviparity and the duration of their foraging trips, seabirds have evolved obligate biparental care during both incubation and chick rearing, which has led to social monogamy (Lack 1968, Jouventin and Cornet 1980, Wittenberger and Tilson 1980, Ligon 1999). An optimal mate choice should enable each mate to assume its parental duties successfully during incubation and chick rearing and to optimize its reproductive output. However, other © 2002 by CRC Press LLC 270 Biology of Marine Birds FIGURE 9.2 Relationships in a subantarctic avian community between foraging range and life history traits such as clutch size (abo ve species name). The key factor in seabirds is the distance between feeding and breeding grounds. Foraging trips, both flying and diving, represent an energetic cost that prevents the most pelagic seabirds from rearing more than one chick per year (or every other year). Are site and mate fidelity a consequence of this low fecundity and high longevity? (Modified from Jouventin and Mougin 1981.) © 2002 by CRC Press LLC Site and Mate Choice in Seabirds: An Evolutionary Approach 271 FIGURE 9.3 Demographic characteristics and reproductive strategy of albatrosses. Delayed maturity results in the presence of high numbers of immature individuals belonging to several age classes. If survival of breeders decreases, immatures recruit into the breeding population at a younge r age than if the population were at equilibrium, acting as a buffer (at least temporarily) against a decline in the breeding population. (Modified from Jouv entin and Weimerskirch 1984b.) © 2002 by CRC Press LLC 272 Biology of Marine Birds constraints than the distance between the breeding grounds and the feeding area may influence the breeding distribution on land and the breeding schedule of seabirds. 9.2.2 PHYLOGENETIC CONSTRAINTS Phylogeny is likely to play a major role in breeding distribution and reproduction in seabirds; however, there are considerable variations of the breeding range within the same group (Figure 9.4; see Chapter 3). Phylogeny also has a strong influence on fecundity. Seabirds lay small clutches (most no more than three eggs; see Appendix 2) and/or have a low fecundity. The 2 Aptenodytes penguins, all procellariiforms, frigatebirds, and tropicbirds, 5 sulids out of 10, 1 gull out of 51, 12 terns out of 44, and 11 alcids out of 22 lay one egg only (see Table 9.2 for within-group variations of clutch size). In addition, many species do not lay replacement clutches (Nelson 1978, Jouventin and Mougin 1981, Warham 1990, Gaston and Jones 1998). Moreover, some breed only every other year (Nelson 1978, Warham 1990, Zotier 1990b). However, some species can raise successfully two broods in succession during the same year (del Hoyo et al. 1992, Williams 1995). Seabirds have extended periods of parental care. Incubation varies between circa 20 days in the smallest terns to 79 days in the largest albatrosses (see Appendix 2). The nestling period (i.e., from hatching until departure from the colony) ranges from 2 days in Synthliboramphus alcids (Gaston and Jones 1998) to almost 1 year in King Penguins (Aptenodytes patagonicus; Marchant and Higgins 1990). Moreover, conditions at sea can affect chick growth, which takes longer in years of poor food availability, due to, e.g., El Niño events (see Schreiber and Schreiber 1993, Red- tailed Tropicbirds). In tropical sulids, frigatebirds, skuas, many gulls, terns, and alcids, parents provide postfledging care (from 1 month in some alcids, Gaston and Jones 1998; to several months in frigatebirds and Abbott’s Booby [Sula abbotti], Nelson 1972, 1976). Conversely, chicks of penguins (del Hoyo et al. 1992), petrels, and albatrosses (Warham 1990), tropicbirds (del Hoyo et al. 1992), gannets (Morus sp., Nelson 1978), and puffins (Fratercula sp., Gaston and Jones 1998) must fend for themselves after leaving their natal colony. High adult life expectancy, however, may enable seabirds to compensate for the long duration of their breeding cycles, low fecundity, and mortality of juveniles at sea (see Introduction). Although adult life expectancy is between 12 and 15 years in many seabird species (Table 9.1), some individuals attain very old ages: Northern Royal Albatross (Diomedea epomophora) 61 years (Robertson 1993, see Appendix 2). Some Emperor Penguins and Snow Petrels that the authors banded as breeders in the mid-1960s at the French station of Dumont d’Urville, Terre Adélie (Antarctica), are still alive at over 35 years of age. 9.2.3 PHENOLOGICAL CONSTRAINTS Food is classically considered the ultimate factor that determines the breeding period in most avian species (Lack 1968, Daan et al. 1988). Because energetic demands of birds are highest during breeding, birds generally breed during periods of highest food availability (Perrins 1970, Martin 1987, Harrison 1990). Marine productivity increases with latitude, but undergoes marked seasonal changes (Nelson 1970, Jouventin and Mougin 1981, Harrison 1990) and breeding synchrony is higher in temperate and polar areas (although some exceptions may occur, Croxall 1984) and chick growth becomes faster as latitude increases (Ashmole 1971, Nelson 1983). Accordingly, we checked for a negative correlation between latitude and the duration of chick growth (i.e., until chicks fledge). For each species, latitude was determined by calculating the average value (accuracy: 1°) between the northernmost and the southernmost locality in its breeding area. We excluded the Emperor Penguin from our analyses because chicks of this species depart to sea at only half of adult body mass (Isenmann 1971). Life history theory predicts that large-sized organisms should have a slower growth than small ones (Stearns 1992); therefore, we divided the chick growth period by adult body mass (after assuming that hatchling body mass was negligible compared to adult body mass). We obtained the amount of time necessary to produce a unit of mass (TUM), which © 2002 by CRC Press LLC [...]... al ( 199 6); 14, Jouventin et al ( 199 9a); 15, Jouventin and Bried (in press); 16, Pianka and Parker ( 197 5); 17, Switzer ( 199 3); 18, Carrick and Ingham ( 196 7); 19, Choudhury ( 199 5); 20, del Hoyo et al ( 199 2); 21, Mougin ( 197 0); 22, Bried et al ( 199 9); 23, Davis ( 198 8); 24, Mougeot et al ( 199 8); 25, Ollason and Dunnet ( 197 8); 26, Ens et al ( 199 5); 27, Bried and Jouventin ( 199 8); 28, Gochfeld ( 198 0); 29, ... ( 199 2), Lequette et al ( 199 5), Burger and Gochfeld ( 199 6), Gochfeld and Burger ( 199 6), Furness ( 199 6), Nettleship ( 199 6), and Zusi ( 199 6) © 2002 by CRC Press LLC 274 FIGURE 9. 4 Continued © 2002 by CRC Press LLC Biology of Marine Birds Site and Mate Choice in Seabirds: An Evolutionary Approach FIGURE 9. 4 Continued © 2002 by CRC Press LLC 275 276 Biology of Marine Birds TABLE 9. 2 Duration of Parental Investment... + + © 2002 by CRC Press LLC Biology of Marine Birds For references, see Nelson 197 8, 198 0, Warham 199 0, del Hoyo et al 199 2, Furness 199 6, Burger and Gochfeld 199 6, Gochfeld and Burger 199 6, Nettleship 199 6, and Gaston and Jones 199 8 Site and Mate Choice in Seabirds: An Evolutionary Approach 281 FIGURE 9. 7 Partitioning of breeding habitats in a seabird community on a sub-Antarctic island: Kelp Gulls... existed in larids (Beer 196 9, Evans 197 0; Figure 9. 9) Individual acoustic recognition has ever since been proved in penguins (Jouventin 198 2, Davis and Speirs 199 0, Jouventin et al 199 9b, Lengagne et al 199 9), procellariiforms (Bretagnolle 199 6, Jouventin et al 199 9a), sulids (Nelson 197 8), and terns (McNicholl 197 5, Møller 198 2) According to Bretagnolle ( 199 6), individual recognition would occur more... tropicbirds, and frigatebirds, 14 cormorants, 6 skuas, 29 gulls, 28 terns, 1 skimmer, and 16 alcids; data in Nelson 197 8, del Hoyo et al 199 2, Lequette et al 199 5, Burger and Gochfeld 199 6, Furness 199 6, Gochfeld and Burger 199 6, Nettleship 199 6, Zusi 199 6) This relationship remained significantly negative if we considered each taxonomic order separately, except for Procellariiformes (Figure 9. 5) However,... production of which requires great amounts of energy, and/or through energy-consuming acoustic and/or visual displays (Andersson 199 4; for seabirds, see Genevois and Bretagnolle 199 4 for the Blue Petrel; Harrison 199 0 for frigatebirds) Most of the costs of obtaining a mate are due to intrasexual competition, which may limit opportunities for mate sampling (Andersson 199 4, Johnstone 199 5, Reynolds 199 6) These... JOUVENTIN 199 8 Why do Lesser Sheathbills Chionis minor switch territory? Journal of Avian Biology 29: 257–265 BRIED, J., AND P JOUVENTIN 199 9 Influence of breeding success on fidelity in long-lived birds: an experimental study Journal of Avian Biology 30: 392 – 398 BROOKE, M DE L 197 8 Some factors affecting the laying date, incubation and breeding success in the Manx Shearwater, Puffinus puffinus Journal of Animal... unattended, as occurs in petrels and albatrosses (Warham 199 0), tropicbirds (Schreiber and Schreiber 199 3), frigatebirds, and some boobies (Nelson 198 0) 9. 3 HABITAT SELECTION 9. 3.1 CHOICE OF THE BREEDING PLACE The ultimate factors that determine choice of the breeding place are food and shelter from predators (Lack 196 8, Nelson 198 0, Warham 199 6) Because seabirds leave their young (and sometimes their eggs)... for offspring fitness, early-hatched chicks having higher postfledging survival and sometimes being more successful upon their first breeding attempt than later-hatched young (Perrins 197 0, Nelson 198 0, Visser and Verboven 199 9) In some seabirds, however, breeding success does not significantly increase with pair experience (Shaw 198 6, Williams and Rodwell 199 2, Bried and Jouventin 199 9), but the costs of. .. ostralegus], Ens et al 199 3), or (2) the cost of mate retention is late breeding or not breeding at © 2002 by CRC Press LLC 290 Biology of Marine Birds all, outweighing all potential benefits (see Davis 198 8 for Adélie Penguins; Olsson 199 8 and Bried et al 199 9 for Aptenodytes penguins) A confounding effect of reproductive performance and individual quality on divorce remains possible, i.e., poor-quality individuals . ( 197 4); 89, Phillips ( 198 7); 90 , Nelson ( 197 8); 91 , Aebischer et al. ( 199 5); 92 , Potts ( 196 9); 93 , Boekelheide and Ainley ( 198 9); 94 , Shaw ( 198 6); 95 , Harris ( 197 9a); 96 , Catry et al. ( 199 7); 97 ,. Is. — 93 .6 0 .93 (14.78) 1,418 96 , 97 , 97 C. s. lönnbergi Iles Kerguelen 98 .3 96 .5 0 .92 5 (13.83) 1,835 19, 19, 19, 19 C. s. lönnbergi Anvers Is. — > 89 0 .95 (20.50) 1,700 98 , 98 , 99 C. maccormicki. Adélie 89. 0 90 .9 0 .91 2 (11.86) 1,405 19, 19, 19, 100 C. maccormicki Anvers Is. — > 85 0 .95 (20.50) 1,200 98 , 98 , 99 C. maccormicki Cape Crozier 87.3 98 .5 0 .93 8 (16.63) 1,300 101, 102, 99 L. (novaehollandiae)