Bruce Bagemihl - Biological Exuberance

39

For sex ratios of various Seals and Sea Lions, see Fay 1982:256 (Walrus); for lunulated and salvin’s antbirds, see Willis, E. O. (1968) “Studies of the Behavior of Lunulated and Salvin’s Antbirds,” Condor 70:128—48; for other antbird species with a “surplus” of unmated males but no homosexual pairs, see Willis, E. O. (1969) “On the Behavior of Five Species of Rhegmatorhina, Ant-Following Antbirds of the Amazon Basin,” Wilson Bulletin 81:363–95.

40

Crab-eating Macaque (Poirier and Smith 1974); Pukeko (Craig 1980:594); Rhesus Macaque (Lindburg 1971:14, 69); Tree Swallow (Stutchbury and Robertson 1985, 1987b); Galah (Rogers and McCulloch 1981:90); Scarlet Ibis (Elbin and Lyles 1994:90–91); Flamingo (King 1994:104–5); Nilgiri Langur (Hohmann 1989:449); Little Egret (M. Fujioka, personal communication); Little Blue Heron (Werschkul 1982:382).

41

Black Stilt (Reed 1993:772); Humboldt Penguin (Scholten 1992:6 and personal communication); Savanna (Yellow) Baboon (Rowell 1967a:16, 22—23 [tables 2, 3]); Mallard Duck (Lebret 1961:108 [table I]).

42

Pig-tailed Macaque (Oi 1990a:340); Bottlenose Dolphin (Wells 1991:222); Cheetah (Eaton and Craig 1973:252); Koala (Smith 1980:184); Canada Goose (Collias and Jahn 1959:484); Flamingo (C. E. King, personal communication); Lesser Flamingo (Alraun and Hewston 1997:175–76).

43

Japanese Macaque (Chapais and Mignault 1991:172; Wolfe 1984:155); Giraffe (Dagg and Foster 1976:28, 124, 144); Greylag Goose (Huber and Martys 1993:160). Likewise, in Northern Fur Seal populations with up to 40 or more females for every male, a number of behavioral and other factors insure that nearly every female is still able to mate heterosexually (Gentry 1998:167, 192–93). For Macaques, some researchers have suggested that females resort to homosexuality when deprived of “novel” male partners rather than of males per se (i.e., when they “run out” of new partners or become overly familiar with them) (Wolfe 1984:155, 1986:274 [Japanese Macaque]; Huynen 1997 [Rhesus Macaque]). As Vasey (1996:550) points out, however, this explanation is flawed because the females they turn to are no more “novel” than the males are (and probably even less so, owing to the high levels of female bonding and familiarity in these species). In addition, some females continue to choose other females as partners even in populations that have novel males.

44

Gorilla (Robbins 1996; Fossey 1983, 1984; Harcourt et al. 1981); Hanuman Langur (Weber and Vogel 1970) Crested Black Macaque (Reed et al. 1997; Dixson 1977); Squirrel Monkey (DuMond 1968; Travis and Holmes 1974; Akers and Conaway 1979; Denniston 1980; Mendoza and Mason 1991); Walrus (Miller and Boness 1983; Sjare and Stirling 1996); Lion (Schaller 1972; Chavan 1981); Mallard Duck (Bossema and Roemers 1985; Schutz 1965:457–59); Black-headed Gull (Kharitonov and Zubakin 1984); West Indian Manatee (Hartman 1971, 1979); Cheetah (Eaton and Craig 1973; Eaton 1974a). In some of these cases (e.g., Gorillas, Hanuman Langurs) homosexual activity among males is much more common in same-sex groups although it still occurs sporadically or “residually” in mixed groups; in other cases (e.g., Squirrel Monkeys, Crested Black Macaques) homosexual activity is equally if not more common in at least some mixed-sex groups.

45

Squirrel Monkey (Talmage-Riggs and Anschel 1973:68, 71); Long-eared Hedgehog (Poduschka 1981:81).

46

Silver Gull (Mills 1991:1523, 1526); Mallard Duck (Schutz 1965:442); Canada Goose (Collias and Jahn 1959:500); Jackdaw (Röell 1979:126, table 1); Lesser Scaup Duck (Bellrose 1976:344); Caribou (Bergerud 1974:432).

47

Flamingo (Wilkinson 1989:53-54; King 1994:105; C. E. King, personal communication); Laughing Gull (Hand 1981:138-39); Humboldt Penguin (Scholten 1992:5); Gentoo Penguin (Stevenson 1983:192); Pied Kingfisher (Moynihan 1990:19; Reyer 80:220); Peach-faced Lovebird (Fischdick et al 1984:314); Galah (Rogers and McCulloch 1981:90); Bicolored Antbird (Willis 1967:112).

48

Cattle Egret (Fujioka 1986b:421-22); emperor and other penguins (Williams, T. D. [1995] The Penguins: Spheniscidae, pp. 80, 160 [Oxford: Oxford University Press]); dipper (Wilson, J. D. [1996] “The Breeding Biology and Population History of the Dipper Cinclus cinclus on a Scottish River System,” Bird Study 43:108—18); Oystercatcher (Heg and van Treuren 1998); Australian noisy miner (Dow, D. D., and M. J. Whitmore [1990] “Noisy Miners: Variations on the Theme of Communality,” in P. B. Stacey and W. D. Koenig, eds., Cooperative Breeding in Birds: Long-Term Studies in Behavior, pp. 559—92 [Cambridge: Cambridge University Press]); spotted sandpiper (Oring, L. W., J. M. Reed, and S. J. Maxson [1994] “Copulation Patterns and Mate Guarding in the Sex-Role Reversed, Polyandrous Spotted Sandpiper, Actitis macularia,” Animal Behavior 47:1065-72).

49

Redshank (Nethersole-Thompson and Nethersole-Thompson 1986:228); mustached warbler (Fessl, B., S. Kleindorfer, and H. Hoi [1996] “Extra Male Parental Behavior: Evidence for an Alternative Mating Strategy in the Moustached Warbler Acrocephalus melanopogon,” Journal of Avian Biology 27:88—91); Ostrich (Bertram 1992:125-26, 178); Greater Rhea (Navarro et al. 1998:117-18); Tree Swallow (Leffelaar and Robertson 1985); tropical house wren (Freed, L. A. [1986] “Territory Takeover and Sexually Selected Infanticide in Tropical House Wrens,” Behavioral Ecology and Sociobiology 19:197-206); barn swallow (Crook, J. R., and W. M. Shields [1985] “Sexually Selected Infanticide by Adult Male Barn Swallows,” Animal Behavior 33:754-61); Black Stilt (Pierce 1996:85); Silver Gull (Mills 1989:388); Herring Gull (Burger and Gochfeld 1981:128); African Elephant (Buss and Smith 1966:385—86; Kühme 1963:117).

50

White-handed Gibbon (Edwards and Todd 1991:234; Reichard 1995 a,b; Mootnick and Baker 1994); Ostrich (Sauer 1972:737); Buff-breasted Sandpiper (Lanctot and Laredo 1994:8; Pruett-Jones 1988:1748).

51

American Bison (Komers et al. 1994:324; D. F. Lott, personal communication); Bonobo (Hashimoto 1997:12— 13).

52

Musk-ox (Smith 1976:37, 56, 75-77; Gray 1979; Reinhardt 1985); Asiatic Elephant (Poole et al. 1997:304, 306—7 [fig. 5]); New Zealand Sea Lion (Marlow 1975:186, 203); Wolf (Zimen 1981:140); Killer Whale (Rose 1992:73, 83—84, 112, 116).

53

Ruff (Hogan-Warburg 1966:178—79, 199-200; van Rhijn 1991:69); Pukeko (Jamieson et al. 1994:271; Jamieson and Craig 1987a); Ocher-bellied Flycatcher (Westcott 1993:450); Ruffed Grouse (Gullion 1981:377, 379—80); Oystercatcher (Heg and van Treueren 1998: 689—90); Brown-headed Cowbird (Rothstein et al. 1986:150, 154—55, 167; Darley 1978); Guianan Cock-of-the-Rock (Trail and Koutnik 1986:209).

54

Giraffe (Dagg and Foster 1976:123; Innis 1958:258-60); Japanese Macaque (Vasey 1996 and personal communication; Corradino 1990:360; Wolfe 1984); Hanuman Langur (Srivastava et al. 1991); Gray Seal (Backhouse 1960:310); Killer Whale (Jacobsen 1990:75-78); Zebras (Rasa and Lloyd 1994:186); Great Cormorant (Kortlandt 1949); Orange-fronted Parakeet (Hardy 1965:152-53); Wapiti (Lieb 1973:61; Graf 1955:73; Harper et al. 1967:37); Ducks (McKinney et al. 1983). Most of these cases are also examples of a “preference” for homosexual activity in the participating individuals.

55

White-fronted Amazon Parrot (Clarke 1982:71); Long-eared Hedgehog (Poduschka 1981:81); Steller’s Sea Eagle (Pringle 1987:104); Barn Owl (Jones 1981:54); Rhesus Macaque (Erwin and Maple 1976:12-13); Crab-eating Macaque (Hamilton 1914:307-8); Bottlenose Dolphin (McBride and Hebb 1948:121); Cheetah (Ruiz-Miranda et al. 1998:7, 12); Black-headed Gull (van Rhijn and Groothuis 1987:142-43; van Rhijn 1985:91-93); Mallard Duck (Schutz 1965:442, 449—50, 460).