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2.4 Spatial Learning, Navigation, and Migration 21
VetBooks.ir nuts (Ottoni and Mannu 2001) and black‐ insects or alleviate itching from insect bites;
toss sand or dirt in threats or play; and
handed spider monkeys also engage in tool
use with detached sticks in self‐directed
bodies for thermoregulation and cooling
behaviours (Lindshield and Rodrigues 2009). siphon mud through their trunk onto their
There are several examples of tool use in (Chevalier‐Skolnikoff and Liska 1993).
free‐ranging non‐primate terrestrial species Chimpanzees used leaves to aid in collect-
that have been documented as well. A classic ing and drinking water (Sousa et al. 2009).
example is the use of stones by Egyptian vul- Boinski (1988) reported observations of a
tures (Neophron percnopterus) to break open white‐faced capuchin using a large tree
the shells of ostrich eggs (van Lawick‐Goodall branch as a club to attack a venomous snake.
and van Lawick‐Goodall 1966). California Many more examples of a variety of tool use
sea otters (Enhydra lutris) position stones in wild animals exist both in the scientific
against another stone or on their chest while literature as well in anecdotal reports.
supine and pound mussels repeatedly against
them in order to open the mussels (Hall and
Schaller 1964). Tool use has also been dem- 2.4 Spatial Learning,
onstrated experimentally in many corvids, Navigation, and Migration
including New Caledonian crows and ravens,
which bend hooks from leaf stems to obtain Another way animals learn from their envi-
larvae from logs (Bluff et al. 2010). ronment is how to move through it. To date,
Although the social behaviour of aquatic spe- most experimental demonstrations of how
cies can be particularly difficult to study in the animals use maps and landmarks have been
wild, there are several reports of tool use in conducted in laboratory settings with rats,
marine mammals and fishes as well. Free‐rang- meadow voles (Gaulin and Fitzgerald 1989),
ing bottlenose dolphins (Tursiops spp.) are par- and insects, and even in species with very
ticularly well‐represented in observations of large home ranges. Interestingly, male
tool use, with sponging being the primary tool meadow voles (which have home ranges up
(Krützen et al. 2005), where a dolphin breaks a to ten times larger than females) were supe-
marine sponge off the seafloor and wears it rior to female voles on spatial learning tasks.
over its closed rostrum to apparently probe
into the substrate for fish. South American 2.4.1 Optimal Foraging
fresh water stingrays use water to extract food
from an experimental test apparatus (Kuba Wild animals of all species need to be able
et al. 2010). It is quite clear that tool use can be to successfully navigate adaptively through
highly adaptive for foraging purposes in many their environment for a variety of reasons,
species of distinct taxonomic groups. but perhaps the most essential reason is to
In addition to different species’ biology locate food. But it is not enough to know
and characteristics, certain features of an simply the location of food; it is also vital to
animal’s habitat can facilitate tool use as be able to obtain food efficiently in the wild.
well, such as the availability of potential One of the best‐known concepts in animal
tools or the presence of barriers to a site behaviour and learning is optimal foraging
with additional resources. Western gorillas theory (OFT), which comes from influences
have been observed to use branches as walk- of ethology, ecology, evolution, psychology,
ing sticks to test water deepness in elephant and economics (Dugatkin 2013). OFT is a
pools. African elephants (Loxodonta afri mathematical theory to predict various
cana) have also been observed to use a vari- aspects of animal foraging behaviour under
ety of tools in the wild; elephants will swat certain conditions (Sih and Christensen
and scratch their bodies with vegetation 2001). Because animals have to make a num-
held in their trunks, possibly to remove ber of decisions about foraging under a given