Applied Animal Behaviour Science 140 (2012) 85–91
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Applied Animal Behaviour Science
jou rnal h om epa ge: www.elsevier.com/locate/applanim
Contrafreeloading in maned wolves: Implications for their
management and welfare
Angélica da Silva Vasconcellos
a,∗
, Cristina Harumi Adania
b
, César Ades
a,1
a
Department of Experimental Psychology, University of São Paulo, Av. Prof. Melo Moraes, 1721, Butantã, CEP: 05508-900, SP, Brazil
b
Associac ¸ ão Mata Ciliar, Av. Emílio Antonon, 1000, Chácara Aeroporto, Jundiaí, CEP: 13212-010, SP, Brazil
a r t i c l e i n f o
Article history:
Accepted 17 April 2012
Available online 12 May 2012
Keywords:
Contrafreeloading
Choice tests
Chrysocyon brachyurus
Preference tests
Welfare
a b s t r a c t
Contrafreeloading occurs when animals spend time and effort to obtain food in the presence
of freely available food. There are several interpretations for such an apparent contradic-
tion to optimal foraging models, with an emphasis either on the need to gather and update
information about the environment or on the value of performing species-typical responses.
Evidence suggests that both gathering information about the environment and the expres-
sion of species-typical behaviour are important for the welfare of captive animals. The aim of
the present study was to assess the existence of contrafreeloading in maned wolves (Chryso-
cyon brachyurus), in a situation where animals could get food directly from a “free” source
and/or search and handle hidden food items, an alternative that requires more effort and
is probably more similar to natural foraging conditions. Eight captive, pair-housed maned
wolves were given weekly choice tests in which they could obtain food either by approach-
ing the usual food tray in one section of the enclosure (Tray), and/or by searching for food
at variable sites amongst the vegetation in the other section of the enclosure (Scattered).
Results indicate that maned wolves spent more time in the Scattered than in the Tray sec-
tion of the enclosure (P = 0.02) and that they obtained about half of the food from that
section (48.54% ± SE 0.69). Our results, the first to demonstrate contrafreeloading in maned
wolves, have implications for the husbandry and welfare of this endangered species.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
Contrafreeloading occurs when an animal chooses to
work for food in the presence of identical and easily obtain-
able food. After it was first described in rats by Jensen
(1963), this apparently paradoxical effect has been demon-
strated in several species (Bean et al., 1999; de Jonge et al.,
2008; Inglis et al., 1997; Lindqvist et al., 2002; McGowan
et al., 2010; Menzel, 1991; Reinhardt, 1994; Rozek and
Milam, 2011). Inglis et al. (1997) have summarised the
major explanations for contrafreeloading: (1) aspects of
∗
Corresponding author. Tel.: +55 11 4368 5143; fax: +55 11 3091 4357.
E-mail addresses: angelv@usp.br, angelicavasconcellos@gmail.com
(A. S. Vasconcellos).
1
Deceased.
the earned food alternative become endowed with sec-
ondary rewarding value sufficient to maintain the costly
performance; (2) neophobic tendencies cause withdrawal
from free food offered in a familiar “working” context; (3)
stimulus change associated with earned food increases its
rewarding value; (4) working for food, especially if this
means the performance of species-typical behaviour, may
be reinforcing in its own right; (5) the Information Primacy
Hypothesis states that contrafreeloading is adaptive in a
natural environment, since it is advantageous for a wild
animal to invest a certain amount of energy searching for
possible future food sites. The energy spent in this activity
would be compensated by the possibility of more efficient
feeding intake in the long run.
Osborne (1977) proposed that feeding activity should
only be considered contrafreeloading when the propor-
tion of food that is difficult to obtain exceeds 50% of the
0168-1591/$ – see front matter © 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.applanim.2012.04.012