Impuls IV Ness in Great Apes

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Dynamic Inconsistency in Great Apes
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Laura Salas-Morellón* Ignacio Palacios-Huerta Josep Call
July 2021
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Abstract
This paper studies the nature of temporal preferences in humans by studying their
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basic structure in our closest living primate relatives. We investigate whether in-
tertemporal choices in three great ape species are dynamically consistent. We present
orangutans (Pongo abelii ), bonobos (Pan paniscus) and gorillas (Gorilla gorilla) with
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several series of choices among different quantities of food available after different de-
lays. When apes can select either a reward that is available immediately or one reward
three times bigger available three minutes later, they tend to act impatiently choosing
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the sooner, smaller reward. They also show high within-group variability. However,
when a six-minute delay is added to both alternatives, apes tend to act patiently
and show a more homogeneous behavior. Additional tests with constant size or delay
suggest that participants trade-off the size and delay of the rewards in a temporal
horizon extended up to several minutes. These results represent the first time that
dynamic inconsistency is documented in our closest living primate relatives. As such,
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they are consistent with the hypothesis that time-inconsistent preferences are part of
our evolutionary heritage.
Keywords: Intertemporal Choice, Discounting, Apes, Cognition.
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JEL Codes: D01, D91.

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* Department of Clinical Psychology and Psychobiology, University of Barcelona, Barcelona, Spain, and
Department of Developmental and Comparative Psychology, Max Planck Institute for Evolutionary An-
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thropology, Leipzig, Germany. Email: laurasalasmo@gmail.com

Department of Management, London School of Economics and Ikerbasque Foundation at UPV/EHU.
Email: i.palacioshuerta@gmail.com
School of Psychology & Neuroscience, University of St Andrews, St Andrews, United Kingdom. Email:
jc276@st-andrews.ac.uk.
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This preprint research paper has not been peer reviewed. Electronic copy available at: https://ssrn.com/abstract=3887401
1 Introduction
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We confront time-sensitive choices in many areas of our lives, including dietary choices,
relationships, education, spending, fertility, crime and financial decisions. Intertemporal
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trade-offs are ubiquitous and as such they represent a major issue of interest in economics,
psychology, neuroscience, biology, sociology and other sciences.1
Time preferences can be formally captured by a discount curve that describes how we
value a good as a function of the time we need to wait to obtain it. As is well known,
the traditional discounted utility model entails discounting the value of a delayed reward
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in an exponential manner (Frederick et al., 2002; Samuelson, 1937). Future benefits are
discounted at a constant rate over time, and this implies that we consider each delay equally,
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irrespectively of its position in our temporal horizon. However, research over the last decades
has shown that our time preferences may be better captured by a hyperbolic discount
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function where the long-term discount rate is smaller than the short-term discount rate.
This readily generates a tendency to make more impatient decisions when a delay is nearer
in time.2 For example, when offered the choice between 100 dollars today or 110 dollars
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in one month many of us would be tempted to accept the immediate reward, but if given
the same choice one year in advance (100 dollars at a given date or 110 dollars one month
after that date) many would likely switch to the second option. Such a preference reversal
suggests that our decisions depend on our relative position in the temporal continuum and
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thus may tend to change over time. This form of dynamic inconsistency has been found to
apply to different types of rewards and timescales (Frederick et al., 2002) and to be present in
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different cultures (Wang et al., 2016).3 Similarly, temporal preferences can also be modeled
as a drive for immediate gratification that can be overridden by cognitive systems of self-
control with some associated utility costs.4 Both types of frameworks have greatly enriched
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1
See Berns, Laibson and Loewentein (2007) and many references therein. Patience, for instance, cor-
relates with a range of measures of adaptation and health (e.g., addiction, obesity) and criminal behavior
(Åkerlund et al. (2016), MacKillop et al. (2011), Schlam et al. (2013)).
2
See, e.g., Laibson (1997), Phelps and Pollak (1968), Marzilli-Ericson et al. (2015), van den Bos and
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McClure (2013).
3
See O’Donoghue and Rabin (1999), Strotz (1955), Thaler (1981), Camerer and Loewenstein (2004),
and Lempert and Phelps (2016). Palacios-Huerta (2003) shows how the study of dynamically inconsistent
preferences dates back at least to Adam Smith and David Hume.
4
See, for example, Benabou and Pycia (2002), Bernheim and Rangel (2004), Fudenberg and Levine
(2006), Gul and Pesendorfer (2001), and Thaler and Shefrin (1981). This alternative models do have
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different policy implications but they are often observationally equivalent and hard to differentiate in most
choice datasets. See Benabou and Pycia (2002), Dekel and Lipman (2012), Toussaert (2018) and Palacios-
Huerta and Volij (2021).
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the study of traditional models of intertemporal decision-making in recent years.
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This paper is concerned with the nature of our temporal preferences. The persistence of
dynamic inconsistencies in humans has led some scientists to think that “the problem may
not come from some extraordinary condition that impairs the normal operation of inten-
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tionality, but rather from the process by which all people, perhaps all organisms, evaluate
future goals” (Ainslie and Haslam, 1992). But this process has yet to be fully understood.5
Specifically, despite the interest in assessing the scope of time inconsistency and the under-
lying mechanisms, its social and evolutionary roots remain unclear in the literature. The
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objective of this study is to contribute to this effort.
In general, the origins of any human behavior can be traced back to an interaction
between the experiences of an individual and an innate default that can be either human-
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specific or shared with other species. Traditionally, the most common way of assessing our
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inherited primate background is to evaluate the behavior of our nearest living relatives (see
Haun et al., 2006 as an example). As indicated by Chen, Lakshminarayanan and Santos
(2006) in their study of different behavioral biases with capuchin monkeys, “it is widely
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accepted in both cognitive science and evolutionary psychology that a mechanism is most
likely evolutionarily ancient if it explains analogous behavior in both humans and primates.
That is, since primates and humans are closely related, it is unlikely that a common trait
evolved in parallel between our two species, and much more likely that common traits
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evolved once during our common evolutionary heritage” (see also Santos and Chen (2009)).
We take this approach in this paper.
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Here, we test for the dynamic consistency of preferences in three species of great apes.
These are cognitively flexible primates which, together with the human genus and several
extinct species, conform the Hominidae family. Non-human primates are particularly useful
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for a number of reasons. They are biologically similar to humans. In the case of the bonobos,
gorillas and orangutans that we study, research suggest that they may share with us up to
99% of the same DNA. They possess physiological, anatomical, and neurological systems
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that are present in humans but are lacking in other animals. They are also capable of
learning complex cognitive tasks that assess brain functions in ways similar to humans and
can engage in complex social interactions. As such, they are particularly interesting and
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have highly distinct advantages relative to other animals for our purposes. In our work,
we will study if apes make more patient decisions when their alternatives are both far in
5
See, e.g., Rubinstein (2003) and Soman et al. (2005).
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time, as opposed to when a set of choices includes one immediate alternative. Although this
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pattern has been found in the preferences of other non-human species,6 it has never been
documented in ape species before.
We position the contributions of our work within existing literatures:
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Firstly, assessing the presence of dynamic inconsistency in closely related species can
help, together with studying children and adults from different cultures (Bettinger and
Slonim, 2007; Wang et al., 2016), to assess how likely is this feature to grow in humans
independently of enculturation processes and to endure regardless of external inputs such
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as education or market discipline.
Secondly, if such a fundamental structure of human decision-making is a part of our
inherited cognitive structure, then the assumption of preference stability over time and
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circumstances made in social sciences would be reinforced (Chen et al. (2006)). Importantly,
we note that preference stability does not imply that preferences are static (Stigler and
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Becker (1977), Becker (1976, 1996), Meier and Sprenger (2015)).
Third, decades of research has repeatedly demonstrated the value of non-human pri-
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mates for research in economics and other social sciences, ranging from studies in game
theory and behavioral biases to research on child development and the study of intergenera-
tional impact of early advantage on health and social status (see Chen, Lakshminarayanan
and Santos (2006), Martin, Bhui, Bossaerts, Matsuzawa and Camerer (2014), Dettmer,
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Heckman, Pantano, Ronda and Suomi (2020), and many references therein). We contribute
to this important literature by studying the nature of intertemporal preferences.
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Fourth, cross-species comparisons can contribute in diverse ways to ongoing debates

about decision-making processes and endogenous preference formation in humans (Santos
and Rosati, 2015). For example, assessing the appearance of a behavior in diverse species
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that possess (or lack) a candidate cognitive trait can help us to trace the abilities that are
necessary for such behavior to emerge, and also to disentangle its underlying psychological
mechanisms which, in the case of dynamic inconsistency, are still unclear (Hayden, 2015;
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Scheres et al., 2013). Moreover, knowing how a “behavioral bias” evolved can inform us
of its adaptive value in certain environments and its biological rationality. This knowledge
6
See Ainslie’s (1974, 1975) early work with pigeons. Later on, ample evidence of dynamic inconsistency
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was found in a collection of experiments with pigeons and rodents (see Rachlin (2009) or Vanderveldt et
al. (2016) for reviews). More recent studies from neurobiology and psychopharmacology also indicate that
the intertemporal choices of rhesus monkeys may be better fit by a curve with a non-constant rate of
time preference (Freeman et al., 2012; Hayden & Platt, 2007; Huskinson et al., 2015; Hwang et al., 2009;
Minamimoto et al., 2009).
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may have a key role in the understanding of our own heuristics and in the development of
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more accurate models of human decision-making (De Petrillo and Rosati, 2019; Hayden,
2019).
Finally, our study makes a methodological contribution of interest in its own right from
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the perspective of the comparative effort to study intertemporal choices in a wide range
of species with regard to their evolutionary heritage. Indeed, non-human animals face
time-sensitive decisions in both social and physical contexts, and their temporal choices
can modelled in the lab.7 These efforts have mainly focused on documenting temporal
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discounting in the short run, that is in studying trade-offs between outcomes in which one of
them is always immediate, not in settings in which both are delayed. The methodology that
researchers have applied to a larger number of species is the “delay-adjusting procedure”
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(Tobin et al., 1996), in which an animal can choose between a quantity of food to be
delivered immediately and a three times bigger quantity that, if chosen, is delivered after a
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delay. This delay is adjusted over trials until it reaches an indifference point in which the
animal chooses equally each alternative. This procedure has revealed significant differences
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in the waiting capacities of different species: while pigeons and rats will typically wait only
a few seconds to get a larger reward, great apes are willing to wait up to an average of one
minute (orangutans, bonobos and gorillas) or two minutes (chimpanzees), with other species
falling somewhere along the continuum (see Stevens and Mühlhoff (2012) for a review).8 In
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general, there is some evidence that species with larger body sizes and larger home ranges
behave more patiently, although the sampling of species is often too small to draw solid
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conclusions.9
In the case of great apes, it is known that they are able to prepare for events occurring
up to hours or days later (e.g., Mulcahy and Call, 2006; van Schaik et al., 2013), and there
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has been considerable recent research interest in their future-oriented behavior (Osvath and
Martin-Ordas, 2014; Redshaw and Bulley, 2018). However, time preference in great apes
has only been studied regarding one immediate and one delayed reward. Studying their
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responses to differentially delayed payoffs, as we do in this study, is our contribution to

7
But see, e.g., Carter et al. (2015) and Hayden (2015).
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Some of the differences among species, both in this procedure and in other time-related tasks, can be
attributed to variations in their ecological niche (Santos and Rosati, 2015). For instance, some features of
each species’ feeding ecology can explain why, despite of their close phylogenetic distance, chimpanzees are
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more patient (and risk-seeking) than bonobos (Rosati et al., 2007) or why common marmosets are more
willing to wait for a reward, but less prone to travel for it, than cotton-top tamarins (Stevens et al., 2005).
9
See Stevens (2014), Stevens and Mühlhoff (2012), and also Amici et al. (2008), and Thom and Clayton
(2014).
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what is an important and sizeable literature from this perspective. Our results show that
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this procedure helps to reach a more complete understanding of their valuation of future
rewards in this and likely other species as well.10
The rest of the paper is structured as follows. Section 2 goes over the experimental
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methodology, including subjects, materials, procedures, design and data scoring. Section 3
presents the results, section 4 a discussion, and section 5 concludes. Complete subject data
are provided in an appendix. All experiments have been video recorded.
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2 Experimental Methodology
We assess whether three great ape species exhibit dynamically consistent preferences by
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evaluating their intertemporal choices in two different time ranges. To that end, we present
orangutans, bonobos and gorillas a standard intertemporal choice task in which they can
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choose either one unit of an appetizing food delivered at a given point in time or three units
of the same food delivered three minutes after that point.
Figure 1 illustrates the options presented to the subjects throughout the different phases
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of the study.
[Figure 1 here]
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In condition 1 (test), one option is available immediately after the choice and the other
one three minutes after. In condition 2 (test), the earlier option is not available until six
minutes have elapsed while the other one requires a nine-minute delay. To the best of our
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knowledge, this condition represents the first time that intertemporal choices between de-
layed outcomes has been studied in our closest living primate relatives (but see Hopper et al.,
2015). As the relative temporal distance between rewards is maintained across conditions,
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no difference should be observed between them under a constant rate of time preference;
however, a more patient behavior would be expected in the second condition if participants
exhibit dynamically inconsistent time preferences with temporal immediacy effects. Besides
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these test conditions, series of choices in which the alternatives differ only in size (pretest)
or in time (controls, see Figure 1) are also administered. Each option we present to the
subjects will be referred to as either A or B throughout this study as defined in Figure 1.
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10
See Amici et al. (2008), Rosati et al. (2007), and also Beran et al. (2014), Dufour et al. (2007),
Latzman et al. (2015), Parrish et al. (2014), and Stevens et al. (2011).
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All participants first complete the pretest and then the experimental conditions (1 and 2)
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in a counterbalanced order.
Subjects: 15 apes socially housed at the Wolfgang Köhler Primate Research Center
in Leipzig, Germany are tested: 6 orangutans (Pongo abelii), 5 bonobos (Pan paniscus),
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and 4 gorillas (Gorilla gorilla). The mean age of the sample is 19 years (ranging from 5
to 29 years) and 67% of them are females. All subjects previously participated in a time
preference study when they completed tasks similar to condition 1 of this experiment, and
have been tested in a variety of cognitive and behavioral tasks throughout their lifetime.
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Participants are never water or food-deprived and can stop participating at any time. Table
1 reports the subject data, including rearing history and order in which they receive the
conditions.
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[Table 1 here]
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Materials: The apparatus consist of a flat sliding platform attached to the lower part
of an upright Plexiglas panel, and two containers filled with food. Three circular holes at
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the base of the Plexiglas panel allow the ape to choose among the containers, as indicated
in Figure 2.
[Figure 2 here]
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All subjects were acquainted with the basic choice procedure before the beginning of the
study. Throughout the experiment, we use five kinds of plastic trays that differ in shape
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and color as containers, and banana slices and grapes as rewards.

Procedure: Apes are tested individually, except for females with dependent offspring.
At the beginning of the trial, the experimenter places two containers on the platform and
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fills them with either one, two or three units of food (see description of Design below).
Next, the experimenter pushes the sliding platform forward, thus allowing the subject to
choose one of the containers by touching it (Figure 2). Then, the experimenter pulls the
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platform backward, moves the unchosen container out of sight and starts the delivery of
food according to a delay associated with the kind of container that has been selected (see
Design). If the chosen container implies an immediate delivery, the experimenter gives its
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contents to the subject right away. If the container is associated to a delayed delivery, the
experimenter leaves the room and comes back in time to give its contents to the subject
according to the specific delay (either 3, 6 or 9 minutes). The selected container and the
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food inside it remains in full sight of the ape for the whole waiting period. Once the ape has
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received the reward, the experimenter gathers up the empty container and the trial ends.
During the pretest, when only immediate rewards are available, identical trays are used
as containers for the whole sample. During condition 1 and condition 2, when the rewards
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are delivered with four different delays (0, 3, 6 or 9 minutes), one distinctive tray (different
to that used in the pretest) is presented for each delay, with the correspondence between
container and delay counterbalanced across the sample of subjects.
The procedure of this study was designed in accordance with the German laws on animal
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experimentation and was approved by the joint ethics committee of the Max Planck Institute
for Evolutionary Anthropology and Zoo Leipzig.
Design: The general procedure involves the presentation of two alternatives for the
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subjects to choose from. Each alternative consists of an amount of food that is available
after a length of time. The precise amount of food and/or the length of time required to

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get it varies as a function of the experimental phase and condition (Figure 1).
Pretest phase: The pretest phase is designed to ensure that subjects prefer three
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units of food over one (Figure 1). All apes complete one series of 12 trials and finish this
phase if they choose option B at least 10 out of 12 trials. The subjects that fail to meet
this criterion complete a second series of 12 trials in which the same criterion is applied. If
they don’t meet the criterion in the second series, they are dropped out of the experiment.
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Control phase (condition 1 & condition 2): The control phase assesses whether the
subjects discriminate between the different temporal delays (Figure 1). In order to allow
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the subjects to learn the temporal contingencies of the task, we introduce forced-choice
trials in which only one of the alternatives is presented and available to choose. All apes
complete one series of 12 forced-choice trials (6 for each alternative) followed by one series
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of 12 free-choice trials. Only the free-choice trials are scored. If the ape chooses option A
in at least 10 out of 12 free-choice trials, he passes directly to the next phase. The apes
that fail to meet this criterion receive an additional set of 12 forced-choice trials and 12
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free-choice trials before they proceed, independently of their results, to the next phase.
Test phase (condition 1 & condition 2): Here, we test for the preferences of the
subjects when facing a decision that implies a trade-off between delay and size of the reward
(Figure 1). As in the control phase, we use forced-choice trials to allow the subjects to
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experience the outcomes of both alternatives before experiencing the free-choice trials. The
test phase of each condition consists for all apes in one series of 12 forced-choice trials (6 for
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each alternative) followed by one series of 12 free-choice trials. Only the free-choice trials
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are scored.
All apes complete the pretest first followed by condition 1 and 2 in a counterbalanced
order. Thus, the sequence they receive was “pretest–condition 1–condition 2” for half of the
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subjects and “pretest–condition 2–condition 1” for the other half. Within both condition 1
and condition 2, all apes complete the control phase first before they receive the test phase.
Each subject is scored in 12 or 24 trials (depending on whether they receive one or two
series) in the pretest, 12 or 24 trials in each control, and 12 trials in each test.
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Two trials per session are run in all phases (unless the subject fails to respond to the
first trial, in which case no further trial is presented) and a maximum of one session per
day. We counterbalance the order of side assignments and trial types (the latter being only
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applicable to forced-choice trials) within each series of trials and each session.
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Data scoring and analysis: The choice for each subject and trial is scored live
and checked later against videotapes by the experimenter. Additionally, we had a second
observer code a random sample of 20% of the trials. He agreed with the experimenter in
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98% of the sampled trials (Cohen’s k = 0.96). We calculate the percentage of choices of
option A and B for each subject within each phase and use the latter as the dependent
measure for statistical testing. Only the data from the animals who complete the whole
procedure are considered for the analysis. Given our sample size, we use non-parametric
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techniques. All reported tests use two-tailed probability tests.

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3 Results
One gorilla (Gorgo) failed to meet the pretest criterion and was excluded from the study.
The analysis of the data of the remaining apes reveals no effect of species in any of the scores
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(Kruskal-Wallis test: pretest: H(2) = 1.50, p > 0.05; control of condition 1: H(2) = 5.93,
p > 0.05; test of condition 1: H(2) = 1.99, p > 0.05; control of condition 2: H(2) = 2.50,
p > 0.05; test of condition 2: H(2) = 0.72, p > 0.05). Also, we do not find any significant
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effect of the order of presentation of conditions 1 and 2 (Mann-Whitney test: control of

condition 1: U = 23.5, N = 14, p > 0.05; test of condition 1: U = 19.5, N = 14, p > 0.05;
control of condition 2: U = 24, N = 14, p > 0.05; test of condition 2: U = 15, N = 14,
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p > 0.05). Therefore, we pool all individuals into a single sample.

Figure 3 presents the median percentage of choices in all phases.
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[Figure 3 here]
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As expected, subjects are sensitive to the delay in the rewards, choosing less frequently
the bigger reward in the test of both conditions than in the pretest (Wilcoxon test: pretest
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vs. test of condition 1: z = −3.17, p < 0.01; pretest vs. test of condition 2: z = −3.30, p <
0.001). Also, within each condition, subjects react to the changes in the magnitude of the
rewards, choosing the more delayed option more frequently in the test phases than in their
respective control phases (Wilcoxon test: condition 1: z = −2.57, p < 0.05; condition 2:
z = −2.95, p < 0.01).
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A comparison between the tests of the two conditions reveals that the apes choose option
B (see Figure 1 for equivalences) more often in condition 2 than in condition 1(Wilcoxon test:
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test of condition 1 vs. test of condition 2: z = −2.14, p < 0.05). With regard to individual
data, half of the subjects show a significant change in the same direction (choosing more
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often option B in the test of condition 2) while one of them (Dunja) displays the opposite
pattern (choosing more often option B in the test of condition 1). A combined analysis
of the individual results indicates that there is a globally significant effect in the main
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direction (Fisher’s method: χ2 (28) = 54.14, p < 0.01), reinforcing the idea that the change
in preferences occurs both at the group and individual levels. In the control phases, the
group results show an analogous effect to that of the test phases, since the apes choose more
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often option B in the control of condition 2 (Wilcoxon test: control of condition 1 vs control
of condition 2: z = −3.01, p < 0.01).
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[Table 2 here]
Next, we examine the preferences within each phase. In the pretest, the group selects
option B above chance (Wilcoxon test: z = 3.33, p < 0.001), as do all the individuals. In the
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test of condition 1, the group does not show a clear preference (Wilcoxon test: z = −1.58,
p > 0.05), although six of the subjects do select option B below chance and one subject
(Dunja) selects it above chance. In the test of condition 2, the group selects option B above
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chance (Wilcoxon test: z = 2.69, p < 0.01), as do two of the subjects. Regarding the data
from the control phases, the analysis reveals that in condition 1, the group selects option B
below chance (Wilcoxon test: z = −3.19, p < 0.01), as well as nine of the fourteen subjects.
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In the control phase of condition 2, the group does not show any preference (Wilcoxon test:
z = −1.09, p > 0.05), although one individual (Dokana) does select option B below chance.
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Therefore, we find a near-equality in the choices of each alternative in the second control
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(Figure 3). Given that delays were signaled by arbitrary cues, in principle this could perhaps
indicate a lack of discrimination between the time magnitudes. In fact, it appears that it
is more difficult for both human and non-human animals to distinguish between longer
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intervals (Allman et al., 2014). However, a complete absence of discrimination does not
seem to explain the increase in patience in the long-run trade-offs for two reasons. Firstly,
it would not account for the test results (considering the pretest data, if they perceived the
delays as equal, they would have chosen almost universally the larger reward). Secondly,
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the participants who select the earlier option above chance (Table 2), thus showing a clear
preference for the six-minute delay (Dokana, over both series of the control, and Padana, in
the last one), fall within the range of the rest of the sample in the test, and near its median.
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Instead, we consider that the evidence from the second control is better explained by a
progressive learning of the temporal contingencies of the task by part of our sample: since,
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within each condition, all subjects complete the control before the test, it is conceivable that
at least some of the apes discriminate better the time intervals when they are presented with
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the test phase.
Further, if we consider more generally the changes across phases reviewed above, we can
observe that in both tests the apes chose option B more often than in the respective control
phases and less often than they do during the pretest. This pattern would be hard to explain
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if the subjects did not react in both conditions to the delay of the rewards as well as to their
size. In order to detect possible differences between conditions in the relative weighting of
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these two parameters, we calculate for each individual the difference scores in each condition
(the difference between the percentage of choices of option B in the control and the test),
finding very similar descriptive statistics in both conditions (in terms of quartiles, condition
1: Q1 = 5.21, Q2 = 16.67, Q3 = 26.04; condition 2: Q1 = 3.12, Q2 = 16.67, Q3 = 25.00)
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and no significant difference between them (Wilcoxon test: condition 1 vs condition 2:

z = −0.118, p > 0.05). Thus, we could not find any change in the data’s structure of the
two conditions. Table 3 reports the correlation between the percentage of choices of option
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B across phases.
[Table 3 here]
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As may be seen, there is no significant relationship at p < 0.05 for any pair of scores.
The only significant correlations at p < 0.10 appear to be within both conditions between
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control and test phases. Therefore, it seems that the best predictor of the apes’ intertemporal
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choices during the test phases, although not significant at conventional levels, would be their
choices in the corresponding control phase (i.e., their reaction to the time intervals). This is
true for both conditions. Also, there is no indication that the strength of this relationship
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differs between conditions.
Overall, we take these results as reinforcing the idea that apes trade off the quantity and
the timing of each reward throughout the whole procedure, including condition 2.
Finally, as we observe that the sample variability is substantially higher in the test of
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condition 1 than in the test of condition 2 (Table 2), we ran paired comparisons of the
variance across control and test phases.11 While there are no significant differences in the
variances neither between each control and its corresponding test phase (Wilcox’s HC4 test:
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condition 1: t(12) = −2.16, p > 0.05; condition 2: t(12) = 0.22, p > 0.05) or between
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controls (t(12) = 1.18, p > 0.05), we find that the apes’ choices do have significantly more
within-group variation in the test of condition 1 than in its analogous test in condition 2
(t(12) = 3.64, p < 0.01).
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4 Discussion
Our findings are consistent with the hypothesis that great apes do not have a constant rate of
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time preference but display dynamically inconsistent preferences with a drive for immediate
gratification. When participants choose between one or three units of food without delay,
they select the latter in a median of 92% of the trials, showing a strong preference for the
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large quantity. However, when they have to wait for three additional minutes to get the
large quantity (while the small one is still immediate), they only choose it in a median of
25% of the trials. Crucially, their preference for the large quantity recovers above chance
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levels (58% of the times) when the small and the large reward are available after six and
nine minutes, respectively. All else equal, great apes significantly tend to make more patient
decisions when facing a choice between future rewards than when facing a choice between
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an immediate and a future reward.

The data on apes’ choices between a small, immediate reward and a large, delayed
reward is consistent with the observation in the literature that in this type of situations
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most participants behave impatiently. In general, waiting for three minutes (relative to an
11
See Wilcox (2015).
12
immediate reward) can be a long time for most species, including humans, when facing a
ed
food-related task. Importantly, however, as we have seen, when a constant delay of six
minutes is added to both options, and apes must wait three additional minutes to get the
larger payoff, they preferably choose to wait. That is, the results indicate that when apes
iew
do not face choices producing an immediate food delivery, which is the standard testing
method in the literature, they are able to make more patient decisions.
Additionally, the difference between the control phases in our study suggests changes
in the differential sensitivity to delay across different time ranges. As the time perception
ev
of humans and non-human animals is non-linear, two points in time are perceived as more
similar as they are farther from the present (Allman et al., 2014). This feature, which has
been proposed to underlie human time inconsistency, could partly explain time preference
r
in non-human species as well (Stephens, 2002). Several proposals have shown that our time
discounting may be closer to an exponential function when calculated over our subjective
er
timeline (e.g., Bradford et al., 2019; Zauberman et al., 2009) and that timing processes
of both in human and non-human subjects correlate with their intertemporal choices (see
pe
Marshall et al. (2014) and Wittmann and Paulus (2008) for reviews). It is conceivable
that, in non-human animals, differences between species in time perception may correlate
with different temporal decisions as well. While many time preference studies in non-human
animals do not include controls for the reaction to time, the evidence and variability across
ot
conditions and individuals that we observe in our results would recommend doing so.
We do not find significant differences between species, though this may be due to a
tn
lack of statistical power. In contrast, however, we do find large individual variation and
changes in the degree of this variability across the different time ranges. We also find large
individual differences when participants make choices that involve an immediate reward,
replicating the finding in the literature that, like humans, non-human animals exhibit ample
rin
individual differences in time preference within the near future. However, the within-group
variability substantially decreases in the condition with two delayed choices. Great apes
prefer the larger reward as a group and show quite a homogeneous behavior. This suggests
ep
that previously documented differences in impulsivity between individuals diminish when

deciding about outcomes that are farther in time. Thus, it is possible that differences in
decisions when an immediate reward is available could be related to the capacity to use self-
Pr
control mechanisms that allow humans and some non-human species, such as chimpanzees,
to overcome temptation (Evans and Beran (2007), Mischel and Mischel (1987)). Moreover,
13
the observed variations in the short-term decisions may be more closely related to the basic
ed
motivational processes that are engaged when immediate rewards are processed, and less
associated to the more cognitive processes that likely prevail when deciding among delayed
outcomes (Kim et al., 2012). When we consider events that are distant in time, a more
iew
abstract construal is elicited and higher-level judgements are produced, which could explain
why decisions affecting our future selves are more linked to long-term goals, less self-centered,
and roughly approximate the choices we would make for another person (Liberman and
Trope, 2003; Pronin et al., 2008). In non-human animals, they could also recruit partially
ev
different mechanisms, especially as the decisions’ elements grow more distant in time or
refer to non-present situations.
The study of the reaction of non-human animals to delayed rewards has significant impli-
r
cations for processes of anticipation and representation (Berns et al., 2007), especially when
they concern decisions between future payoffs. Many species can anticipate and mentally
er
represent future events (Osvath, 2015; Raby and Clayton, 2009; Suddendorf and Corballis,
2007) and it has been suggested that future-oriented cognition is related to future-oriented
pe
choices. Human vivid mental representations of future scenarios covary with more patient
decisions (Bulley et al., 2016) and one of the proposed functions of the human capacity of
mental time travel is to facilitate patient and flexible intertemporal choices (Boyer, 2008;
Bulley et al., 2016). Further, the processing of temporally distant payoffs is believed to
ot
overlap with that of socially distant or hypothetical outcomes and to be linked to abstract
construal (Liberman and Trope, 2003; Trope and Liberman, 2010). Therefore, studying the
tn
covariation of these abilities across living species could help us establish their evolutionary
history and adaptive impact. Additionally, intertemporal choices may be a direct means
to explore future-oriented cognition (Osvath and Osvath, 2008). Under natural conditions,
delayed payoffs tend to have minimal perceptual cues and a patient response often requires
rin
some capacity to foresee future outcomes. Although this is true when an individual makes
a patient choice between an immediate-present and a delayed-absent reward (but see Red-
shaw and Bulley, 2018), we agree with Cheke, Thom, and Clayton (2011) that an in-depth
ep
study of the choices between delayed and invisible rewards (e.g., by looking for effects that
are observable with immediate and/or visible rewards) could enrich the possibilities of this
approach.
Pr
Finally, previous work has revealed an intertwinement between time preferences and the
different adaptive strategies of different species, and there is wide agreement that patience
14
may have played a key role in the evolution of the human lineage by enabling a wide
ed
range of critical behaviors, such as cooperation or tool use (Stevens and Hauser, 2004;
Warneken and Rosati, 2015). However, organisms continually face intertemporal decisions
and their available courses of action only sometimes produce an immediate outcome. For
iew
example, when an animal can exploit different patches of food, his set of alternatives only
includes an instant payoff whenever he is already in one of the patches; instead, it will often
choose among options with various non-null temporal costs. We argue that cross-species
comparisons applied to choices between delayed rewards may reveal different patterns that
ev
complement those found in the literature. As such, it would be beneficial to consider
choices between outcomes with different future delays, together with sets of alternatives that
include an immediate benefit, as is typically done, in order to gain a deeper understanding
r
of the different decision-making mechanisms across species and the evolutionary history of
patience.
5 Conclusions
er
pe
The importance of understanding intertemporal trade-offs for many real world phenomena
has led to a long-standing interest in modeling intertemporal decision processes, especially
in the past two decades. Intertemporal choices may vary across goods, from activity to
activity, culture to culture, and person to person. And yet, despite the fact that the economic
ot
literature studying these trade-offs, including temptation, self-control and the demand for
commitment, is voluminous, it includes little work that studies the nature of intertemporal
tn
decision making.
We have approached this question taking advantage of evidence in the evolutionary
sciences showing that if two closely related primates species show analogous behavior, then
rin
it is widely accepted as the most likely explanation that this trait is caused by mechanisms
that are evolutionary ancient and appeared in a common antecessor. We consider that
the study of great apes, with whom we may share up to 99% of the same DNA, is a
ep
main contribution of our study. Together with the study of children and adults across
different cultures, the evidence from our closest related species and other species supports
the hypothesis that time-inconsistent preferences are part of our evolutionary heritage, and
Pr
hence suggests that this feature is likely to grow independently of enculturation processes
and to endure regardless of external inputs such as education or market discipline.
A second contribution is that the analysis builds for future empirical research by sug-
15
gesting that experimental methods with great apes may also be used to distinguish the
ed
origins of dynamic inconsistency and costly self-control, both when preferences are deter-
ministic and stochastic. Although this would indeed be quite challenging, these are two
leading formal frameworks developed in the broad literature that studies temptation and so
iew
worth considering. Studying apes’ behavior could similarly be useful to study the nature
of temptation mechanisms involving visceral influences and cue-contingent influences (see
Berns et al., 2007).
Finally, our findings indicate that direct cross-species comparisons that include choices
ev
between two or more delayed rewards have the potential to explain better the observed
decisions of non-human animals in natural settings, and hence can be useful to provide a
more comprehensive approach to the study of their time preferences. We argue that focusing
r
on intertemporal choices between future events involving rewards that are not present at
the time of the decision constitute a valuable approach that may improve our study of
future-oriented cognition.
er
pe
ot
tn
rin
ep
Pr
16
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25
ed
FIGURE 1 – EXPERIMENTAL OPTIONS AND CONDITIONS
Options presented to the subjects throughout the different phases of the
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study. Number of dots represents the quantity of food units. The time lapse
below each option indicates the delay of the food. A and B labels denote the
short names used in this study.
v
re
er
pe
ot
tn
rin
ep
Pr
ed
FIGURE 2 – EXPERIMENTAL SETUP
iew
At the beginning of the trial, the platform is in the backward position (left
figure). Later, the experimenter pushes the platform forward, thus allowing
the subject to touch one of the containers (right figure).
v
re
er
pe
ot
tn
rin
ep
Pr
ed
FIGURE 3 – APES’ CHOICES
Median percentage of choices of each alternative as a function of study

phase. Error bars indicate semi-interquartile range.
v iew
re
er
pe
ot
tn
rin
ep
Pr
ed
TABLE 1 – AGE, GENDER, REARING HISTORTY
AND OTHER CHARACTERISTICS
iew
Order in
Previous
which
participation
they
Species Subject Age (years) Sex Rearing in temporal
received
preferences
condition
studies
1 and 2
Orangutan Kila 8 Female Mother a 2-1
v
(Pongo 1-2
Padana 10 Female Mother a
re
abelii)
Dokana 19 Female Mother a 1-2
Pini 20 Female Mother a 2-1
Bimbo 28 Male Nursery a 2-1
Dunja 35
er Female Mother a 1-2
Bonobo Yasa 11 Female Mother b 2-1
pe
(Pan 2-1
Kuno 12 Male Nursery b
paniscus)
Limbuko 13 Male Nursery b 1-2
Ulindi 15 Female Mother b 1-2
Joey 26 Male Nursery b 2-1

ot
Gorilla Kibara 5 Female Mother a 1-2
(Gorilla 2-1
Viringika 13 Female Mother a
tn
gorilla)
Gorgo 27 Male Nursery a *
Bebe 29 Female Mother a 1-2
Notes: Previous participation in temporal preferences studies is denoted as: a = Amici

rin
F, Aureli F, and Call J (2008). “Fission-fusion dynamics, behavioral flexibility, and inhibitory
control in primates.” Current Biology 18(18): 1415-1419, and b = Rosati AG, Stevens JR, Hare
B, and Hauser M (2007). “The evolutionary origins of human patience: Temporal
preferences in chimpanzees, bonobos, and human adults.” Current Biology 17(19): 1663–
ep
1668. The right-most column denotes the order of presentation of the two experimental
conditions for each subject. The asterisk * denotes the fact that Gorgo did not receive any
of the conditions as he failed to reach the pretest criterion (see Design and Results).
Pr
ed
TABLE 2 – PERCENTAGE OF CHOICES OF OPTION B IN EACH FASE
AND BARNARD’S PROPORTION TEST BY SUBJECT
iew
Condition 1 Condition 2
Comparison between tests
Subject Pretest Control Test Control Test
KilO 87.5 *** 12.5 *** 8.3 ** 41.7 83.3 * z=3.69 p = 0.000
PadO 83.3 * 0.0 *** 8.3 ** 29.2 50.0 z=2.24 p = 0.029
v
DokO 100.0 *** 0.0 *** 16.7 * 25.0 * 58.3 z=2.11 p = 0.045
re
PinO 100.0 *** 16.7 ** 0.0 *** 33.3 50.0 z=2.83 p = 0.007
BimO 83.3 * 62.5 75.0 45.8 66.7 z=-0.45 p = 0.780
DunO 87.5 *** 33.3 100.0 *** 62.5 58.3 z=-2.51 p = 0.014a
YasB 91.7 ** 8.3 ** er 25.0 58.3 83.3 * z=2.87 p = 0.007
KunB 100.0 *** 0.0 *** 16.7 * 37.5 50.0 z=1.73 p = 0.106
LimB 100.0 *** 12.5 *** 0.0 *** 66.7 75.0 z=3.79 p = 0.000
UliB 100.0 *** 0.0 *** 58.3 45.8 50.0 z=0.41 p = 0.839
pe
JoeB 83.3 * 0.0 *** 25.0 50.0 75.0 z=2.45 p = 0.023
KibG 91.67 ** 45.8 75.0 58.3 58.3 z=0.87 p = 0.525
VirG 100.00 *** 33.3 50.0 58.3 58.3 z=0.41 p = 0.839
BebG 91.67 ** 33.3 50.0 33.3 50.0 z=0.00 p = 1.000

ot
Median 91.67 12.5 25 45.83 58.33

(IQR) (13.54) (33.33) (54.17) (25) (25)
tn
Notes: This table reports the percentage of choices of option B in each phase, and the
results of Barnard’s exact test for the comparison of two proportions for each subject. The
percentage of trials in which the ape selects alternative B is calculated for each subject and
rin
phase as the number of choices divided by the number of completed trials (12 or 24).
Species membership is indicated by a superscript on the subject’s name (O = orangutan, B
= bonobo, G = gorilla). Asterisks next to percentages denote a significant deviation from
chance according to a binomial test: * = p < 0.05; ** = p < 0.01; *** = p < 0.001. Upperscript
“a” denotes a significant change in the opposite direction from that of the hypothesis, and
ep
IQR is the inter-quantile range. The raw number of choices is available in the appendix.
Pr
ed
TABLE 3 – SPEARMAN’S RANK CORRELATION COEFFICIENTS BETWEEN
THE PERCENTAGE OF CHOICES OF OPTION B ACROSS PHASES
iew
Pretest Control Test Control Test
v
rs = 1.00 rs = -0.164 rs = -0.272 rs = -0.016 rs = -0.325
Pretest
p = 0.575 p = 0.346 p = 0.956 p = 0.257
re
rs = 1.00 rs = 0.492 rs = 0.382 rs = 0.129
Control
p = 0.074* p = 0.177 p = 0.660
Condition 1
rs = 1.00 rs = 0.371 rs = -0.029
Test
p = 0.192 p = 0.922
er rs = 1.00 rs = 0.532
Control
p = 0.050*
Condition 2
pe
rs = 1.00
Test
Note: * denotes significant at 10% level.

ot
tn
rin
ep
Pr
ed
APPENDIX - Choice data
iew
Number of choices of option B within each series of trials for each subject.
v
Pretest Control Test Control Test
Subject
re
1st 2nd 1st 2nd 1st 2nd
series series series series series series
Pad 10 0 1 5 2 6
Dok 12 0 2 3 3 7
Dun 9 12 4 er 4 12 8 7 7
Bim 10 8 7 9 5 6 8
Pin 12 4 0 0 3 5 6
Kil 9 12 3 0 1 5 5 10
Uli 12 0 7 5 6 6
pe
Kun 12 0 2 4 5 6
Lim 12 3 0 0 10 6 9
Yas 11 1 3 5 9 10
Joe 10 0 3 5 7 9
Beb 11 4 4 6 3 5 6
Kib 11 5 6 9 7 7 7
ot
Vir 12 3 5 6 6 8 7
tn
rin
ep
Pr

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Dynamic Inconsistency in Great Apes

JEL Codes: D01, D91.

thropology, Leipzig, Germany. Email: laurasalasmo@gmail.com

Fourth, cross-species comparisons can contribute in diverse ways to ongoing debates

responses to differentially delayed payoffs, as we do in this study, is our contribution to

and color as containers, and banana slices and grapes as rewards.

techniques. All reported tests use two-tailed probability tests.

effect of the order of presentation of conditions 1 and 2 (Mann-Whitney test: control of

p > 0.05). Therefore, we pool all individuals into a single sample.

and no significant difference between them (Wilcoxon test: condition 1 vs condition 2:

an immediate and a future reward.

that previously documented differences in impulsivity between individuals diminish when

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Options presented to the subjects throughout the different phases of the

Median percentage of choices of each alternative as a function of study

Dokana 19 Female Mother a 1-2

Pini 20 Female Mother a 2-1

Bimbo 28 Male Nursery a 2-1

Limbuko 13 Male Nursery b 1-2

Ulindi 15 Female Mother b 1-2

Joey 26 Male Nursery b 2-1

Gorilla Kibara 5 Female Mother a 1-2

Gorgo 27 Male Nursery a *

Bebe 29 Female Mother a 1-2

Notes: Previous participation in temporal preferences studies is denoted as: a = Amici

PadO 83.3 * 0.0 * 8.3 29.2 50.0 z=2.24 p = 0.029

YasB 91.7 8.3 er 25.0 58.3 83.3 * z=2.87 p = 0.007