an adult (Whitehead <strong>and</strong> Weilgart 1991). When a predator is encountered (such as a killer whale),the group adopts what is called the marguerite or wagon-wheel formation (Whitehead 2002a). Thisformation consists of females in a circle, aligned like the spokes of a wheel, with their tail flukesforming the rim <strong>and</strong> the calves in the centre, or hub, of the wheel (Whitehead 2002a). The femalesput themselves at risk to protect the other members of the group, in particular wounded animals<strong>and</strong> calves. The whales will slash their tail flukes at predators (including humans, whom theyperceive as predators) who try to attack individuals in the group. An alternative protectivearrangement is the ‘heads out’ formation wherein the animals arrange themselves in a tight rank,with their heads (<strong>and</strong>, therefore, their teeth) facing towards the predator, with calves, again, in thecentre of the formation (Whitehead 2002a).Although these defensive formations assist the sperm whale to defend itself against naturalpredators, such as sharks <strong>and</strong> killer whales, humans unfortunately have used the whale’s defensivebehaviour to their advantage during whaling activities. <strong>Whale</strong>rs have killed group members one byone, knowing that their fellows would st<strong>and</strong> by <strong>and</strong> not leave the sides of injured or strickenanimals (Tyack 2002a). Exploiting the bond between mother cetaceans <strong>and</strong> calves, whalers in StVincent <strong>and</strong> the Grenadines have traditionally targeted a humpback calf in order to lure its mothercloser to the whaling boat (see chapter 6).St<strong>and</strong>ing by injured group members is a behaviour that has been observed in many other cetaceanspecies, particularly when attempting to keep a stricken animal’s blowhole above the sea surface inorder that it can still breathe (Caldwell <strong>and</strong> Caldwell 1966). There have been many records ofcetaceans supporting dead companions, especially calves, long after the animals have died,sometimes for a period of several days. This type of behaviour has been reported in a variety ofspecies, including common dolphins (Dephinus delphis), rough-toothed dolphins (Steno bredanensis),striped dolphins (Stenella coeruleoalba), Pacific humpback dolphins (Sousa chinensis) <strong>and</strong> bottlenosedolphins (Tursiops truncatus) (Moore 1955; Brown et al. 1966; Lodi 1992; Fertl <strong>and</strong> Schiro 1994;Parsons 1998).WHALES – INDIVIDUALS, SOCIETIES AND CULTURES19These collaborative behaviours reflect not only intelligence, but also show that individuals withingroups benefit from the presence of the other group members. In addition, individuals may havediffering, but complementary, roles. Thus, the removal of any one animal may negatively affect theremainder.SocietiesA society is defined as “an interacting group consisting of more individuals than parents <strong>and</strong> theirimmediate dependent offspring” (Slobodchikoff <strong>and</strong> Shields 1988). Several species of cetacean formmatrilineal societies, the best-studied being sperm whales, pilot whales (Globicephala spp.) <strong>and</strong> somekiller whales. Within these societies, females spend their entire lives with their natal group(families). Male killer whales, <strong>and</strong> possibly male short-finned pilot whales (G. macrorhynchus), staywith these female groups (that is, they stay with their mothers, sisters, <strong>and</strong> aunts), but mate withunrelated females from other groups they encounter (thus avoiding inbreeding).The matrilineal groups of pilot whales <strong>and</strong> killer whales are particularly interesting; female pilotwhales <strong>and</strong> killer whales may live 20 years beyond the end of their reproductive years (that is, theyexperience menopause), living past 60 years of age. It has been suggested that this allows old females
to store <strong>and</strong> pass on information to other group members <strong>and</strong> perhaps provide alloparental care inthe form of babysitting <strong>and</strong>/or wet nursing (Heimlich-Boran <strong>and</strong> Heimlich-Boran 1994; Acevedo-Gutiérrez 2002; Baird 2002). Pilot whales (<strong>and</strong> also sperm whales) suckle calves for 13 to 15 years,indicating a very long period of dependency <strong>and</strong> close association with their mothers.Sperm whale females may also form associations that last for decades. These groups appear to assistin the care of young animals, with group members taking the role of babysitters, to allow mothersto take long, deep dives in search of food (Whitehead <strong>and</strong> Arnbom 1987; Whitehead et al. 1991).It has been suggested that the cultural transmission of learned behaviours to family members isconserved in matrilineal groups: these societies assist in the learning of complex skills essential forsurvival, as well as ensure these skills are passed onto future generations without being lost orforgotten. Although these societies aid in the transmission of information, they also have the effectof changing mating patterns <strong>and</strong> hence the genetic evolution of the populations <strong>and</strong>/or species(Whitehead 1998). Genetic modification as the result of ‘culture’, such as occurs in these cetaceansocieties, was previously believed to occur only in humans (Acevedo-Gutiérrez 2002).20A REVIEW OF THE WELFARE IMPLICATIONS OF MODERN WHALING ACTIVITIESCultureCulture can be defined as behavioural variations between sets of animals that are maintained <strong>and</strong>transmitted by social learning (Whitehead 2002b) <strong>and</strong> typically involves components of bothteaching <strong>and</strong> imitation by the animals concerned. It has been suggested that everything ofimportance in human behaviour is transmitted culturally (Manning <strong>and</strong> Dawkins 1992). Culturehas been identified in several cetacean species, for example, in the eastern North Pacific, killer whalegroups were discovered to possess distinct calls that are unique to their group members (Ford 1989,1991, 2002). In short, like many populations of humans, these cetaceans had unique vocal dialects.Similar dialects have also been found in other species of cetacean (Ford 2002), including the codasof sperm whales (Weilgart <strong>and</strong> Whitehead 1997) <strong>and</strong> the songs of humpback whales (Payne <strong>and</strong>Guinee 1983). These dialects are learned from older members of the group in the matrilineal spermwhale <strong>and</strong> killer whale, <strong>and</strong> from other singing males on the breeding grounds of humpback whales;that is, cultural transmission of information rather than ecological or genetic differences account forthe geographical differences in vocalisations (Rendell <strong>and</strong> Whitehead 2001).Rendell <strong>and</strong> Whitehead (2001) identified several different ways in which culture was transmitted incetaceans. The first was a spread of novel <strong>and</strong> complex behaviours between members of the samegeneration (i.e. a ‘horizontal’ transmission of culture), which occurs in humpback <strong>and</strong> bowheadwhales when learning songs from other males (e.g. Noad et al. 2000). The second type of culturaltransmission was between mother <strong>and</strong> young (i.e. a ‘vertical’ transmission of culture); for example,female killer whales teaching offspring how to str<strong>and</strong> themselves in order to catch the pups ofsouthern sea lions (Otaria flavescens) or elephant seals (Mirounga leonina) on haul-out beaches(Lopez <strong>and</strong> Lopez 1985; Guinet <strong>and</strong> Bouvier 1995). The final pattern of cultural transmission hasbeen described above; a stable group transmission of culture that can be horizontal, vertical or‘oblique’ (a non-parent from a previous generation transferring information), e.g. vocal dialects inkiller whales (Deecke et al. 2000) or sperm whales (Weilgart <strong>and</strong> Whitehead 1997).Other apparently culturally transmitted behaviours include bottlenose dolphins placing sponges ontheir beaks to avoid being punctured by urchins <strong>and</strong> spiny fish when foraging; bottlenose dolphins,
- Page 5 and 6: ForewordWhales are highly evolved a
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for the task. The correct target ar
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11 Review of criteria for determini
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It is apparent from Figure 1, that,
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Table 1 Animal welfare and the Sche
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16 Summary of conclusionsModern day
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Glossary136A REVIEW OF THE WELFARE
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Appendix IIColour plates©Mark Voti
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142A REVIEW OF THE WELFARE IMPLICAT
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Figure 13. Processing minke whales