Black-Capped Chickadees (Poecile atricapillus) Anticipate Future
Outcomes of Foraging Choices
Miranda C. Feeney, William A. Roberts, and David F. Sherry
University of Western Ontario
In 2 experiments we investigated the cognitive abilities of wild-caught black-capped chickadees (Poecile
atricapillus) in future anticipation tasks. Chickadees were sensitive to anticipatory contrast effects over
time horizons of 5, 10, and 30 min (Experiment 1). Chickadees also learned the order of events and
anticipated that the quality of future foraging outcomes was contingent on current foraging choices. This
behavior was demonstrated while foraging in a naturalistic aviary environment with a 30-min delay
between the initial choice and the future outcome (Experiment 2). These results support the hypothesis
that black-capped chickadees can cognitively travel in time both retrospectively and prospectively using
episodic memory. This result shows the occurrence of anticipatory cognition in a noncorvid species of
food-storing bird and supports the idea that cognitive time travel may have evolved in nonhuman animals
in response to specific ecological selection pressures.
Keywords: future anticipation, mental time travel, anticipatory contrast, food storing
Within the declarative memory system, memory for personally
experienced events, episodic memory, is distinguished from mem-
ory for facts about the world, semantic memory (Tulving, 1972).
Mental time travel (MTT) is the ability to cognitively re-
experience past events or to plan for future events and depends on
both episodic and semantic memory (Schacter, Addis, & Buckner,
2008; Suddendorf & Corballis, 2007). Episodic memory in hu-
mans includes the property of autonoetic consciousness for per-
sonally experienced past events. The fundamental distinction of
episodic memory is that it allows people to remember what, where,
and when events happened along with a sense of reliving the
episode (Tulving, 1985, 2002). Although Tulving (1983) argued
that episodic memory is a uniquely human ability, there has been
debate over whether nonhuman animals have a form of episodic
memory. Although autonoetic consciousness cannot be accessed in
animals, a number of recent experiments suggest that some ani-
mals can remember what, where, and when an event occurred. This
ability has been called what-where-when (WWW) memory or
episodic-like memory (Babb & Crystal, 2005; Clayton & Dickin-
son, 1998; de Kort, Dickinson, & Clayton, 2005; Roberts, 2002,
2006; Roberts & Feeney, 2009; Suddendorf & Corballis, 1997,
2007; Zinkivskay, Nazir & Smulders, 2009).
There has been ongoing research evaluating the episodic-like
memory capabilities of nonhuman animals over the last decade
(Babb & Crystal, 2005; Bird, Roberts, Abroms, Kit, & Crupi,
2003; Clayton & Dickinson, 1998, 1999b; Feeney, Roberts, &
Sherry, 2009; Hampton, Hampstead, & Murray, 2005; Zinkivskay
et al., 2009). More recently, theorists have begun to examine the
importance of the episodic and semantic memory systems for
future anticipation and planning (Atance & O’Neill, 2001; Buck-
ner & Carroll, 2006; Correia, Dickinson, & Clayton, 2007; Raby,
Alexis, Dickingon, & Clayton, 2007). Information stored in epi-
sodic and semantic memory may be used to generate elaborate
mental scenarios describing the anticipated future, suggesting that
MTT is a complex, constructive ability. Suddendorf and Corballis
(2007) argued that the ability to use knowledge of past experiences
to adaptively plan for the future provided the primary selection
pressure for the evolution of brain structures responsible for hu-
man MTT. Schacter et al. (2008) subsequently advanced the con-
structive episodic simulation hypothesis, proposing that episodic
memory and future simulation are based on common neural pro-
cesses.
In nonhuman animals, Eichenbaum and Fortin (2009) demon-
strated that rats can flexibly integrate memories to predict out-
comes in novel situations. They argued based on neuropsycholog-
ical and behavioral evidence that animals have the capacity for
retrospective recollection and future prediction and that both abil-
ities rely on the hippocampus in nonhuman animals, as is the case
for human MTT. Although animal MTT may not be as elaborative
or constructive as human MTT, the similarities in neural correlates
suggest that animal MTT may be relatively complex in its own
right. Raby and Clayton (2009) argued that even if animal MTT is
qualitatively different from that of humans, the possibility for
semantic prospection—thoughts about the future without concur-
rent self-projections—is an overlooked area of research that has
been hindered by a focus on human MTT.
Flaherty and Checke (1982) foreshadowed the recent interest in
future planning and anticipation with an examination of anticipa-
tory contrast, or the suppressed intake of a lesser value food at
Time 1 in favor of a higher value food at Time 2. Results showed
This article was published Online First August 16, 2010.
Miranda C. Feeney, William A. Roberts, and David F. Sherry, Depart-
ment of Psychology, University of Western Ontario.
Support for this research was provided by Discovery Grants from the
Canadian Natural Sciences and Engineering Research Council to William
A. Roberts and David F. Sherry.
Correspondence concerning this article should be addressed to Miranda
C. Feeney, Department of Psychology, University of Western Ontario,
London, Ontario N6A 5C2 Canada. E-mail: mcfeeney@uwo.ca
Journal of Experimental Psychology: © 2010 American Psychological Association
Animal Behavior Processes
2011, Vol. 37, No. 1, 30 – 40
0097-7403/10/$12.00 DOI: 10.1037/a0019908
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