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18  2  The Cognitive Abilities of Wild Animals

  VetBooks.ir  reinforcement that would be likely to occur   intrasexual competitions by bighorn sheep
                                                     do not always result in a won contest for that
            in free‐roaming animals, it can sometimes be
            difficult to determine whether a behaviour is
            increasing due to positive reinforcement or   individual animal.
            negative reinforcement. If we observe a griz-  2.2.2  Punishment
            zly bear swimming in a river on a hot sum-
            mer day, one might wonder whether the bear   Just like both positive and negative rein-
            is swimming because the behaviour (i.e.   forcement, animals experience punishment
            swimming) is providing coolness or opportu-  in many situations in their natural environ-
            nities for foraging (and is thus positively   ment. Within social groups of primates, for
            reinforcing) or because the behaviour is   example, subordinate animals who attempt
            resulting in the removal of discomfort from   to access preferred resources (e.g. food,
            overheating  (and  thus negatively reinforc-  reproduction  with potential  mates) may
            ing)? In a mobbing situation, do European   often be punished by dominant members of
            bee‐eaters exhibit aggression toward snakes   the group. Another example can be thought
            because mobbing results in the removal   of in terms of mobbing  –  an antipredator
            of an aversive predator near their eggs (i.e.   behaviour  –  in birds toward other avian
            negatively reinforcing) or because mobbing   predators. The function of mobbing behav-
            provides increased opportunities for social   iour is described as a means to bring
            cohesion among bee‐eaters or a higher num-  together conspecifics  –  usually within a
            ber of intact eggs (i.e. positively reinforcing).   social group  –  to remove intruders (Caro
            Although these are empirical questions that   2005). A hawk or eagle that is mobbed by a
            require further scientific study to tease apart,   flock  of crows  might be less  likely to  visit
            it is nonetheless interesting to consider exactly   that spot because it resulted in the aversive
            what processes are contributing to the learn-  consequence of being attacked by multiple
            ing history that modifies an individual’s   blackbirds simultaneously (i.e. positive pun-
            behaviour over its lifetime.             ishment) or losing a preferred food item in
              In addition to different types of reinforce-  the process (i.e. negative punishment).
            ment, the schedule at which an animal may   Individuals who leave a food source
            obtain reinforcers in the wild varies tremen-  unguarded without caching or storing their
            dously. Broadly speaking, animals may obtain   food, may find that as a consequence, other
            reinforcers  continuously  or  intermittently.   individuals – either from within or outside a
            An animal that is receiving continuous rein-  social unit – can run off with these resources.
            forcement for a behaviour is obtaining rein-  This would be a form of negative punish-
            forcement every time it exhibits that    ment,  in which  the removal  of  appetitive
            behaviour, whereas behaviours that are inter-  stimuli (i.e. resources) can follow from
            mittently reinforced are not reinforced every   overly aggressive or passive behaviour.
            time. Interestingly, behaviours that are  not   Scientific evidence of negative punishment
            reinforced every time are actually more likely   is much less prevalent than positive punish-
            to persist longer in the absence of reinforce-  ment in wild animals. However, there are
            ment than behaviours that are reinforced   some extreme examples that can be seen in
            every time. As you might suspect, most   cases of infant mortality and disappearances
            behaviours that an individual animal exhibits   in male takeovers in free‐ranging howling
            in the wild are intermittently reinforced. For   monkeys (Alouatta palliata) in Costa Rica
            example, stalking or hunting by a male jaguar   (Clarke 1983), a species in which dominant
            does not always result in successfully taking   males have exclusive access to mate with
            down prey, male peacocks that engage in   receptive  and high‐estrous females. Thus,
            sexual displays in soliciting for a potential   punishment contingencies, although not
            mate do not always achieve copulation, and   strictly programmed as in a captive setting,
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