ORIGINAL PAPER

Social learning in captive African elephants(Loxodonta africana africana)

Brian J. Greco • Tracey K. Brown •

Jeff R. M. Andrews • Ronald R. Swaisgood •

Nancy G. Caine

Received: 1 May 2012 / Revised: 29 November 2012 / Accepted: 30 November 2012 / Published online: 14 December 2012

� Springer-Verlag Berlin Heidelberg 2012

Abstract Social learning is a more efficient method of

information acquisition and application than trial and error

learning and is prevalent across a variety of animal taxa.

Social learning is assumed to be important for elephants,

but evidence in support of that claim is mostly anecdotal.

Using a herd of six adult female African bush elephants

(Loxodonta africana africana) at the San Diego Zoo’s

Safari Park, we evaluated whether viewing a conspecific’s

interactions facilitated learning of a novel task. The tasks

used feeding apparatus that could be solved in one of two

distinct ways. Contrary to our hypothesis, the method the

demonstrating animal used did not predict the method used

by the observer. However, we did find evidence of social

learning: After watching the model, subjects spent a greater

percentage of their time interacting with the apparatus than

they did in unmodeled trials. These results suggest that the

demonstrations of a model may increase the motivation of

elephants to explore novel foraging tasks.

Keywords Elephants � Loxodonta � Social learning �Imitation � Animal cognition

Introduction

Social learning occurs when an animal acquires informa-

tion from another individual’s behavior and then applies

that information in a similar situation at a later time to

achieve a similar objective (Coussi-Korbel and Fragaszy

1995). Psychologists typically categorize social learning

into a variety of imitative and non-imitative forms. For

example, in true imitative social learning the observer

copies some portions of a behavioral motor sequence

(Dawson and Foss 1965; Whiten et al. 1996; Zentall et al.

1996; Zentall 2004). The various forms of non-imitative

social learning can take place when the model’s behavior

enhances or gives new value to novel stimuli, objects,

locations, or events (Heyes 1993; Call and Carpenter 2002;

Zentall 2006). For example, social facilitation occurs when

a conspecific’s behavior motivates the observer to engage

in a similar behavior (e.g., foraging-specific behaviors)

(Fragaszy and Visalberghi 1990; Zentall 2006). Stimulus

enhancement plays a role in non-imitative social learning

when the actions of a model draw the attention of an

observing animal toward a specific stimulus or part of a

stimulus (e.g., an object or location) (Call and Carpenter

2002; Whiten et al. 2004; Zentall 2006). Observational

conditioning allows an observing animal to learn some-

thing about the relationship between two stimuli, as occurs

when an observer learns from watching a model that a food

All experimental procedures were approved by the San Diego Zoo’s

IACUC #09-014.

Electronic supplementary material The online version of thisarticle (doi:10.1007/s10071-012-0586-7) contains supplementarymaterial, which is available to authorized users.

Present Address:B. J. Greco (&)

Department of Animal Science, One Shields

Avenue, Davis, CA 95616, USA

e-mail: bjgreco@ucdavis.edu

B. J. Greco � J. R. M. Andrews � R. R. Swaisgood

San Diego Zoo Global, San Diego, CA, USA

T. K. Brown

Department of Biological Sciences, California State University

San Marcos, San Marcos, CA, USA

N. G. Caine

Department of Psychology, California State University San

Marcos, San Marcos, CA, USA

123

Anim Cogn (2013) 16:459–469

DOI 10.1007/s10071-012-0586-7

hopper delivers (i.e., is associated with) food (Call and

Carpenter 2002; Zentall 2006). Imitative and non-imitative

learning can and often do co-occur, and it can be difficult to

determine which is operating at a given time (Zentall

2006).

Many social learning experiments rely on an observer–

demonstrator paradigm, which compares the performance

of a non-exposed control group to that of subjects who are

first allowed to observe a conspecific demonstrate a novel

task (Voelkl and Huber 2000; Horner et al. 2006; Dindo

et al. 2008). Following a predetermined number of suc-

cessful demonstrations, the demonstrating animal is

removed from the enclosure, and the observer is allowed to

participate in the experimental task by itself (Voelkl and

Huber 2000; Horner et al. 2006; Dindo et al. 2008). If the

performance of the observing subjects is significantly dif-

ferent from that of the non-exposed subjects, inferences can

be made about the presence of social learning (Voelkl and

Huber 2000). For example, should the observers reliably

show a bias toward the motor sequence and solution

method demonstrated by the model (i.e., copying fidelity),

imitation may be responsible (Voelkl and Huber 2000;

Caldwell and Whiten 2004; Horner et al. 2006; Dindo et al.

2008). Additionally, changes in performance speed, accu-

racy, and/or the ways in which observing subjects manip-

ulate an apparatus may be interpreted as indicators of

non-imitative social learning (Fragaszy and Visalberghi

1990; Caldwell and Whiten 2004). For example, in a

number of studies of social learning in capuchins (Cebus

apella), Fragaszy and Visalberghi (1990) found that the

behavior of models who solved novel foraging tasks

influenced the actions of group members who had not yet

learned the task on their own. The authors concluded that

increased apparatus exploration time, elevated apparatus

manipulation, and faster solution times by observers were

likely attributable to stimulus enhancement and/or social

facilitation. Evidence of learning by imitation has been

claimed for certain species of birds (Dawson and Foss

1965; Zentall 2004), primates (Whiten et al. 2004), and

cetaceans (Rendell and Whitehead 2001; Krutzen et al.

2005). Non-imitative social learning has been demon-

strated in a larger number of animal taxa, including fish

(Schuster et al. 2006), rats (Ray and Heyes 2002), bats

(Page and Ryan 2006), gray squirrels (Hopewell et al.

2010), ravens (Stowe et al. 2006), horses (Krueger and

Heinze 2008; Krueger et al. 2011), swine (Oostindjer et al.

2011), parrots (Huber and Gajdon 2006), and capuchins

(Dindo et al. 2009), to name just a few.

Female elephants (Loxodonta and Elephas) live in

hierarchical matrilineal family groups composed of 5–10

related adult females and their immature offspring (Moss

1988; Sukumar 2003; Byrne et al. 2009). In contrast,

mature male elephants spend most of their time living

solitarily, only occasionally associating with female herds

or bachelor herds (Moss 1988; Sukumar 2003). In African

bush elephants (Loxodonta africana africana), family

groups are also known to aggregate into large bond groups,

in which the elephants travel, forage, and communally

raise/protect their young (Moss 1988; Sukumar 2003; By-

rne et al. 2009). Their cooperative behaviors are mediated

by acoustic, chemical, seismic, tactile, and visual com-

munication signals (Langbauer 2000).

Given the social cohesiveness of elephant herds and the

strength of social bonds among elephants, there seems to be

ample opportunity for social learning to occur. Surpris-

ingly, however, evidence of social learning in elephants is

limited primarily to anecdotes (Byrne et al. 2009). The

reasons for this curious absence are probably related to the

challenges elephants pose as experimental subjects. Not

only does their extraordinary size make them difficult and

potentially dangerous to manipulate, but constructing

experimental apparatus that can withstand an elephant’s

strength and curiosity is formidable. Furthermore, it is

impossible to maintain the amount of experimental control

that can be achieved with smaller, more tractable animals

such as birds [e.g., Campbell et al. (1999)]. Nonetheless, to

better understand the ways in which elephants adapt and

respond to ecological and social challenges, we need to find

ways to investigate their learning styles and capabilities.

Here we report the results of a controlled study of social

learning in a group of African bush elephants (Loxodonta

africana africana) housed in captivity. We expected that

adult female elephants would use the same technique for

solving a foraging task as a conspecific model. We also

developed two metrics, focus score and initial interest, to

investigate other forms of social learning as well.

Methods

Subjects

The subjects for this study were six female African bush

elephants (Loxodonta africana africana), ranging in age

from 18 to 21 years and living at San Diego Zoo’s Safari

Park. These females are believed to be related, but their

complete histories are not known. The elephants were

wild-born in South Africa and brought to the Safari Park

in 2003 after it was learned that they were to be culled for

the purpose of population control (Andrews et al. 2005).

At the time of our study, all but one of the females had at

least one calf that was born while living at the Safari

Park. In addition to these six subjects, two adult male

African elephants were used to test the safety, function-

ality, and difficulty of the experimental apparatus in pilot

trials.

460 Anim Cogn (2013) 16:459–469

123

The six female subjects and 11 other herd members (two

adult bulls and nine calves) lived in a 2.23-ha outdoor

enclosure. Experimental trials were conducted in two

adjacent, 0.20-ha enclosures separated by a steel-cable

barrier fence through which the elephants could see each

other. During experimental trials, subjects were removed

from the main group for no more than 60 min at a time.

Water was provided during each trial.

Experimental apparatus

The six apparatus used in this study were individually

designed and constructed to be unique and novel to the

subjects. All apparatus delivered food reward (alfalfa-

based dry herbivore pellet or pressed alfalfa) when solved,

but no two apparatus shared the same mechanisms. One

apparatus could only be solved in one way, but the

remaining five apparatus had two distinct solutions that

involved different behavioral actions. Additionally, many

of the apparatus were constructed with either noticeably

movable or breakaway components on the apparatus. Thus,

a subject could cue on the model’s movements, apparatus

movements, the location of apparatus components, and/or

breakaway apparatus components when observing the

model (see Fig. 1 and the supplementary material for

drawings and photographs).

The Kerplunk (K) was a roughly cylindrical apparatus

composed of six open-ended pipes attached to a central

shaft. Four of the six outer pipes were stuffed with pressed

alfalfa bundled in paper (to ensure the alfalfa slid out when

the K was solved). When hung vertically, four bamboo

shafts transected the four stuffed pipes, preventing the

alfalfa from sliding out and preventing the subjects from

pulling the alfalfa out of the apparatus. The K was the only

one of the six apparatus that did not include two different

solution options. In order for a subject to solve the appa-

ratus, she needed to remove two bamboo shafts.

The Counterweighted Feeder (CW) was composed of a

long rope slung over a tree. One end of the rope was free

and accessible to the subjects, while the other end was

weighted at ground level. A bundle of pressed alfalfa was

tied to the free end of the rope out of the subjects’ reach.

When given access to the remaining length of free rope, a

subject could draw the alfalfa bundle toward herself by

pulling. If the rope was not secured, the weight would tug

the alfalfa back out of reach. However, one of two

behavioral motor sequences—(a) securing the slack with

the mouth or (b) securing the slack with a forefoot—could

be used to ‘‘choke-up’’ on the rope and draw the alfalfa

bundle into trunk’s reach.

The Pop and Roll (PR) apparatus was a tube with an

internal trolley-mounted feeder ball. When presented to the

elephants, the PR was hung horizontally, just below the

subjects’ eye level. The elephants were prevented from

freely accessing the feeder ball by two end caps. One end

cap was mounted to the trolley, and the other was a free-

floating plug. To solve the PR, the elephants needed to

either remove the free-floating plug (using an attached

handle) or push on the trolley-mounted cap (which would

force the free-floating plug out of the opposite end of the

tube). From the perspective of an observing subject, the

model solving the apparatus by end cap removal would

present a yanking behavior with her trunk from the left side

of the PR, followed by the emergence of her trunk

clutching the end plug. If the model chose to push the

trolley-mounted end cap, she would present a trunk-

Fig. 1 Pop and Roll a subject solving the apparatus by pulling the

end plug. Notice the outward arch of the trunk as the subject pulls the

end plug free. b Subject solving the apparatus by pushing the feeder

ball to force the end plug out. Notice the trunk is forcefully inserted

into the right side of the apparatus, thus popping the free-floating end

plug out of the left side. For detailed descriptions and photographs of

all apparatus, please see the supplementary material section on

experimental apparatus

Anim Cogn (2013) 16:459–469 461

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thrusting motion from the right side of the PR and the cap

would fall out of the left side. In both scenarios, the model

would retrieve a reward from the left side of the PR. If the

model pushed the trolley-mounted end cap, the subject

might additionally observe the movement of externalized

trolley components.

The Push Pop (PP) was an apparatus with an internal-

ized plunger. When presented to the subjects, the PP was

hung vertically with the plunger arm protruding from the

bottom. Pressed alfalfa was loaded into a cavity on top of

the plunger pad. Subjects could manipulate the plunger by

grasping and thrusting the plunger arm or by striking the

base of the arm with the dorsal surface of the trunk. Both

thrusting and striking caused the plunger to rise, forcing the

alfalfa reward out of the cylinder.

The Boxall (BX) was constructed by combining a plastic

enrichment box and a plastic enrichment ball. The ball was

suspended in the box with one hemisphere protruding from

one of the box’s faces. Two holes were cut into the inner

hemisphere of the ball, and the outer hemisphere was left

intact. On an adjacent face of the cube, a hinged flap was

attached to cover a trunk-sized hole. The apparatus was

presented to the subjects with the ball facing outward and

the flap on the top face of the box. Subjects could solve the

apparatus by rotating the ball, so that there was one hole in

each hemisphere, inner and outer, or by opening the hinged

flap outward. Either action would allow the subject to

insert her trunk into the box to retrieve the alfalfa reward.

The Smash Box (SB) was constructed from a plastic

enrichment box with a weight attached to the bottom face.

The box was modified to pivot end-over-end when

manipulated or struck. On the front face of the SB, a balsa

wood panel covered a 20-cm-diameter hole, and on the top

face of the SB, a rope bung was inserted to plug a 5-cm-

diameter hole. Subjects could solve the SB either by

shattering the balsa wood panel with their tusk or by

pulling out the rope bung. Once the wood was broken or

the bung removed, the box could be spun end-over-end to

collect the food reward.

Procedures

There were two types of trials: modeled, in which a subject

watched the dominant elephant (Swazi; see below) suc-

cessfully operate the apparatus before being allowed to

interact with the apparatus herself, and unmodeled, in

which a subject interacted with the apparatus without

having watched Swazi. Each of the subjects interacted with

three of the apparatus in the modeled condition and three of

the apparatus in the unmodeled condition.

Pilot data from the bull elephants were used to roughly

categorize the six apparatus into easy (K and PP), medium

(CW and SB), and hard (PR and BX) levels of difficulty.

We then attempted to shuffle the order of experimental

trials to allow each subject access to an apparatus of easy,

medium, and hard difficulty in both the modeled and un-

modeled condition, while simultaneously preventing each

subject from acting in the same experimental condition

(modeled or unmodeled) more than two consecutive times.

Additionally, when assigning subjects to condition, we

avoided duplicating any one set of three apparatus (e.g., in

the modeled condition only one subject saw the Counter-

weighted Feeder, the Pop and Roll, and the Smash Box).

Subjects participated in trials about once a week (except for

Swazi, who participated in her own trial and then acted as

model in other trials). One of us (BG) conducted all of the

trials with the aid of an assistant. All trials were videotaped

for later analysis.

Experimental trials: unmodeled condition

Prior to each trial, the apparatus to be tested was mounted

in place. BG called the subject over to the apparatus and

drew her attention to it. He then stepped away, giving the

subject up to 45 min to interact with and solve the appa-

ratus. Once the subject solved the apparatus, she was

allowed to collect any reinforcement distributed by the

apparatus and was given additional food reward to rein-

force her participation. Subjects who failed to solve the

apparatus were also given a food reward at the end of the

45-min trial period, to reinforce participation.

Experimental trials: modeled condition

The most dominant female in the herd, Swazi, served as the

model in all of the modeled trials. Out of view of the rest of

the herd, she was given access to an apparatus in an un-

modeled trial. During this trial Swazi was allowed to solve

each apparatus and was given time to practice that solution

several times. Swazi’s first interaction with each apparatus

was used as data for the analysis of responses in the un-

modeled condition. The decision to use just one model

animal was based on two factors. First, little is known

about elephant social learning and how individual differ-

ences among elephants may influence whether, when, and

how one elephant learns from another. Using just one

model for all subjects controlled for at least one aspect of

the social learning situation (i.e., the identity of the model)

in this study. Second, social hierarchy plays an important

role in elephant herds; if the roles were reversed, a sub-

dominant female may not feel comfortable using an

apparatus that is within close proximity to a more dominant

member of the herd. For this reason, the most dominant

female was selected as the model.

During modeled trials, the apparatus was mounted in

place, and Swazi (the model) was allowed to solve the

462 Anim Cogn (2013) 16:459–469

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apparatus, while the subject was present in an adjacent

enclosure separated by a fence. Swazi always used the

same solution option as she used on her first successful

trial. Once Swazi successfully solved the apparatus, the

apparatus was reset and the observation process was

repeated. A subject was allowed to view Swazi demon-

strate each apparatus two or three times. The number of

demonstrations was kept constant for each individual

apparatus.

Following her demonstrations, Swazi was removed, the

apparatus reset, and the subject given access to the appa-

ratus as if she were participating in an unmodeled trial.

Approximately 5 min passed between Swazi’s demonstra-

tion and observer participation in the modeled trial.

Dependent variables and data coding

Two assistants, blind to condition and subject, indepen-

dently viewed all of the videotaped trials. The assistants

were trained by BG using videos from pilot trials that made

use of apparatus that were not used in the experiment. The

assistants each scored all 30 trials. Fifteen of these sessions

were unmodeled, and fifteen were modeled. Swazi’s un-

modeled sessions also were scored, but these data were not

used to calculate any of the results.

The times scored by the two assistants were averaged for

use in all of the relevant analyses described below. Inter-

rater reliability was high (r [ 0.90) across scorers.

Interaction time was scored from the unmodeled trials

and modeled trials. Because the subjects sometimes turned

their attention away from the apparatus or got distracted

with something unrelated to the apparatus, interaction time

included only those times when the elephant was actually

touching the apparatus in some way. A touch was defined

as any contact with the apparatus that appeared to be

intentional (i.e., trunk, rostrum, foot, or body contact nec-

essary to manipulate an apparatus, but not incidental con-

tact associated with proximity). Solution time is the

duration of interaction time on a particular trial. If an

individual subject solved an apparatus, her solution time

and interaction time would be the same.

Modeled trials were scored for the behavior of both the

model and the observer. First, the assistants scored Swazi’s

interaction time to give us one measure of how much

information was available to the observing elephant in

modeled trials. This is referred to as demonstration time for

a particular trial. Then the interaction time of the subjects

was scored, as above.

In addition to the scoring by the research assistants, BG

also viewed all of the videotapes to extract the following

information: (a) Exposure time: The total amount of time

elapsed between a subject’s first touch of an apparatus and

the solve event. (b) Focus score: Each subject’s interaction

times were divided by her appropriate exposure times to

calculate a focus score. When interaction and exposure

times are similar (a focus score approaching 1.0), it means

that an elephant was very engaged in the task. A low focus

score means that the subject often did not focus her

attention on the task (a focus score greater than 1.0 is not

possible, as interaction time cannot be greater than expo-

sure time). (c) Solution method: which of the two possible

solutions was chosen by the subjects in each trial.

(d) Number of touches: the number of times a subject

intentionally touched an apparatus. High rater reliability

was reached for number of touches. (e) Bouts to solution:

number of bouts each subject required to solve each

apparatus. A bout was defined as any string of continuous

touches occurring within 30 s of one another. Note that a

bout could consist of just one touch or up to 49 touches (the

maximum number of touches recorded per bout). High

rater reliability was reached for bouts to solution. As an

assessment of his reliability and consistency in scoring, BG

scored and later rescored a subset of the trials and achieved

nearly perfect agreement (r = 0.99) on both touches and

bouts. (f) Initial interest: The duration of each subject’s

first bout on an apparatus was divided by her total inter-

action time with that apparatus. Subjects that showed

maximum interest in an apparatus upon first given access to

it have initial interest scores approaching 1.0.

Finally, BG examined the video data from the three

apparatus that featured two separate solution mechanisms

(the PR, BX, and SB) in order to determine whether the

observers touched the apparatus in locations that the model

also touched. This was done by mapping the location of

touches onto schematic drawings of the apparatus. Using

these touch maps, we could estimate the number of times an

observing subject’s touches overlapped Swazi’s touches.

Results

Trials

Copying fidelity

Contrary to our hypothesis, subjects in the modeled con-

dition did not regularly copy the methods used by the

model to solve the novel apparatus. In the thirteen modeled

trials with the apparatus that offered two solutions, four

trials were solved using the same method as the model, six

were solved using the different method, and in three trials

the subject failed to solve the apparatus at all (Figs. 2, 3). If

the initial difficulty ratings are accounted for, solution

failures occurred on two difficult apparatus, one moderate

apparatus, and one easy apparatus. The solutions used by

the subjects in the modeled and unmodeled conditions were

Anim Cogn (2013) 16:459–469 463

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distributed fairly evenly between the two available options

(Fig. 3).

Solution time

The subjects solved the apparatus on 25 of the 30 trials.

Three of the five failures were in the modeled condition.

Ndlula failed to solve apparatus 4, 5, and 6, Umoya failed

at solving the third apparatus, and Litsemba failed to solve

the sixth apparatus. When an elephant failed to solve a trial

within the time limit (45 min), its solution time was

recorded as the highest solution time scored by a subject on

that particular apparatus.

Within-subject analysis revealed no significant differences

for average solution time between the modeled (M = 189.07,

SD = 63.69) and unmodeled conditions (M = 184.93,

SD = 67.81) (paired samples t test: t(4) = 0.08, N = 5,

P = 0.94) (Fig. 4). We hypothesized that subjects would

score better (i.e., lower average solution times) in the modeled

condition than in the unmodeled condition. For one of the

subjects, this was true. However, two of the remaining sub-

jects showed similar average solution times in the modeled

and unmodeled condition, and two showed lower average

solution times than unmodeled subjects.

Focus scores

On average, when subjects participated in the modeled

condition, they displayed focus scores (interaction time

divided by exposure time) (M = 0.46, SD = 0.08) that

were 50.2 % greater than the focus scores they achieved in

unmodeled trials (M = 0.31, SD = 0.08) (paired samples

t test: t(4) = 3.71, N = 5, P = 0.02) (Fig. 5). Indeed, all

five subjects had higher focus scores in the modeled than in

the unmodeled condition. It is interesting to note that those

individuals who failed to solve an apparatus often dis-

played lower focus scores for that apparatus.

Touches and bouts

There was no significant difference in the average number

of touches when subjects were in the modeled (M = 53.7,

SD = 25.9) versus unmodeled conditions (M = 51.6,

SD = 28.6) (paired samples t test: t(4) = 0.2, N = 5,

P = 0.9) (Fig. 6), nor was there a significant difference

in the average number of bouts in the two conditions

(modeled: M = 10.8, SD = 7.3; unmodeled: M = 9.1,

SD = 4.1) (paired samples t test: t(4) = 0.6, N = 5,

P = 0.6).

Touch-map analysis revealed that Swazi touched the

apparatus in solution-relevant and non-solution-relevant

locations during all demonstrations. The total number of

touches ranged greatly, from 10 to 51, and touches directed

toward solution-relevant locations were always weighted

toward the solution mechanism she demonstrated (average

ratio: Pop and Roll = 2:1; Boxall = 21:1; Smash

Box = 5:2). In contrast, the solution-relevant touches of

the modeled condition subjects were more evenly distrib-

uted (average ratio: Pop and Roll = 17:11; Boxall = 19:7;

Smash Box = 7:8) (Fig. 7). On average, 49.8 % of subject

touches overlapped the locations touched by Swazi (the

model) during demonstration, and of these touches, an

average of 25.3 % overlapped solution-relevant locations

and an average of 22.2 % overlapped non-solution-relevant

areas. Additionally, an average of 58.3 % of touches

occurred in areas of the apparatus that had small openings

to the food chamber (e.g., seams, cracks, and tie-down

eyelets), and an average of 37.9 % of these touches over-

lapped areas touched by Swazi.

Initial interest

On average, when subjects participated in the modeled

condition (M = 0.48, SD = 0.22), their initial interest in

an apparatus (interaction time of the first bout divided

by total interaction time) was 102 % greater than the

initial interest of subjects who participated in the unmod-

eled condition (M = 0.24, SD = 0.10) (paired samples

t test: t(4) = 3.11, N = 5, P = 0.036) (Fig. 8). As was

the case with focus scores, all subjects displayed higher

initial interest in the modeled than in the unmodeled

condition.

Umoya

Ndlula

Umngani

Litsemba

Lungile

SF

SF

SF

Fig. 2 The copying fidelity of observing subjects according to

apparatus. Apparatus are represented by the icons from left to rightCounterweighted Feeder, Pop and Roll, Push Pop, Boxall, and SmashBox. Stars indicate a match in solution method. Crossed out boxesindicate solution methods used other than the model’s. Blank cellsindicate that the subject interacted with an apparatus in the

unmodeled condition. SF stands for solution failure. Only two

subjects observed the model demonstrate the Push Pop and the Boxall

464 Anim Cogn (2013) 16:459–469

123

Additional analyses

Concordant with the similar solution times across the two

(modeled and unmodeled) conditions, Swazi’s average

demonstration times were not correlated with the observ-

ers’ average solution times (Pearson’s r = 0.15, N = 5,

P = 0.81). These correlations were calculated again to

compare data from apparatus that had been demonstrated

twice by the model, and then again for data demonstrated

three times by the model. In comparing these correlations,

there was no indication that extra demonstration time

influenced subjects’ performance in the modeled condition.

Discussion

We predicted that the elephants would demonstrate social

learning by using the same technique to solve a foraging

task as they had seen in a demonstration session by a

familiar conspecific. However, the subjects were just as

likely to use an alternate solution as they were to copy the

Mouth Pull ekirtStooF llaBhsuP Thrust Flap Rope Panel

SF: 1

1

3

2

SF: 1 SF: 1

SF: 1

Mod

eled

Unm

odel

ed

1

2

ALT:1

3

SF: 1

Fig. 3 This figure demonstrates

the use of solution options by

apparatus in both modeled and

unmodeled conditions

(including the options Swazi

learned first during model

training). Apparatus are

represented by the icons from

left to right: CounterweightedFeeder, Pop and Roll, PushPop, Boxall, and Smash Box. In

each case the total adds up to six

trials. SF stands for solution

failure. One subject solved the

Pop and Roll via an alternate

method (ALT). Only two

subjects observed the model

demonstrate the Push Pop and

the Boxall

Fig. 4 Average solution times per subject are listed according to

dominance rank, with the most dominant on the left and least on the

right

Fig. 5 Focus score are shown in unmodeled and modeled conditions

Anim Cogn (2013) 16:459–469 465

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actions of the model. Copying fidelity is a measure of true

imitation, whereby an observer uses only the actions of the

model, and not changes associated with the apparatus itself,

to guide its subsequent interactions with that apparatus.

Had we seen evidence of copying fidelity, it would not

have been certain that the criteria for true imitation had

been met, insofar as our apparatus, with the possible

exception of the PP and CW apparatus, were not true ‘‘two-

action’’ apparatus (Dawson and Foss 1965), which are

designed to test for true imitation (Voelkl and Huber 2000).

However, the data from our two (possible) two-action tasks

are not more suggestive of true imitation than the data from

the other tasks; in each case, half of the subjects did not use

the solution method they had seen in the demonstration by

the model.

In addition to copying fidelity, we examined various

measures of performance that can shed light on non-imi-

tative forms of social learning. For example, if subjects

solve a problem more quickly after witnessing a model

successfully gain food reinforcement from an apparatus, it

could indicate that the observation of a model has increased

the motivation of the observers or drawn attention to the

aspects of the apparatus that are relevant to a solution

(Fragaszy and Visalberghi 1990). However, when com-

paring subjects’ scores across the two conditions, the

solution times of the subjects were not significantly dif-

ferent. Furthermore, on three occasions the elephants failed

to solve the apparatus at all, even though they had the

benefit of watching the model successfully solve the

apparatus two or three times in a row.

The ways and extent to which a subject physically

interacts with an apparatus (number/location of touches

and numbers/duration of bouts of touches in the current

study) also can help to elucidate forms and mechanisms of

both imitative and non-imitative learning in a social

learning situation. For example, despite the fact that none

of their subjects was able to solve a novel ‘‘two-action’’

Fig. 6 The mean number of touches are diagrammed in modeled and

unmodeled conditions

26Swazi44Umoya

03Swazi01Lungile

46Swazi42Litsemba

016Swazi38Umngani

226Swazi411Lungile

05Swazi136Umoya

23Swazi30Umngani

68Swazi628Lungile

SR2SR1Model IDSR2SR1Modeled Condition Subject ID

Fig. 7 The number of touches to solution-relevant components of the

Pop and Roll, Boxall, and Smash Box are demonstrated above.

Modeled condition subject touches are tallied on the left, with touches

demonstrated by the model (Swazi) tallied on the right. Each of the

examined apparatus featured two solution-relevant components (SR):

Pop and Roll SR1 = pull plug; SR2 = push trolley; Boxall:

SR1 = ball; SR2 = flap; Smash Box: SR1 = pull rope; SR2 = break

panel

466 Anim Cogn (2013) 16:459–469

123

foraging apparatus, Caldwell and Whiten (2004) found that

common marmosets (Callithrix jacchus) that had observed

a model’s demonstration more frequently touched the

apparatus in locations used by the model than marmosets

that had not seen a model’s demonstration. Such demon-

stration-consistent touching could arise from stimulus

enhancement or social facilitation, both of which are forms

of non-imitative social learning.

In contrast to the findings of Caldwell and Whiten

(2004), the present study failed to reveal evidence of

demonstration-consistent touching by the subjects. Despite

the fact that Swazi’s demonstrated touches were always

weighted (at least 2:1, and usually much greater) toward

the solution mechanism she demonstrated, the attention of

the observing subjects was more evenly distributed. We did

find that an average of 49.8 % of all touches by modeled

condition subjects overlapped locations where Swazi had

touched during demonstration. However, 37.9 % of these

touches overlapped in areas of the apparatus that had small

openings to the food chamber (e.g., seams, cracks, and tie-

down eyelets). Therefore, we believe that the overlap of

subjects’ touches with those of the model is more consis-

tent with an interest in food-based odors emanating from

the apparatus than they are suggestive of learning from the

model.

We created two measures, the focus score and the initial

interest score, to investigate the effects of modeling that

were not addressed by any other variables alone. Focus

scores—a measure of the proportion of time elephants

spent interacting with the apparatus—address a subject’s

engagement in the task at hand. Focus scores were, on

average, 50.2 % higher in modeled trials than in unmod-

eled trials, and all five elephants had higher focus scores in

the former. Initial interest—a measure of the proportion of

interaction time used in the first bout—is similar to that of

focus scores in that it addresses a subject’s engagement in a

task, but it may be a more accurate measure of the

immediate affects of observing a modeled demonstration.

On average, subjects in the modeled condition demon-

strated 102 % more initial interest in apparatus than sub-

jects in the unmodeled condition. Together, these results

point to the possibility that observing the demonstrations of

a model may increase an elephant’s general motivation to

approach and engage a novel foraging task. Interestingly,

the observing subjects in our study showed higher average

focus scores and initial interest despite the fact that they

did not show higher average numbers of touches.

Thus, it seems possible that observational conditioning

may explain the difference in focus scores and initial

interest across conditions. Observational conditioning may

have occurred because the subjects learned that the appa-

ratus could be made to deliver food, motivating them to

interact with the apparatus with the expectation that they

could be rewarded with pellets or hay. Given the social and

spatial cohesiveness of elephant groups, there is ample

opportunity for elephants to benefit from simply directing

their attention to the consequences and contexts of a con-

specific’s actions.

However, we must also consider that when interacting

with the apparatus during a modeling trial, the model

(Swazi) may have left odors on the apparatus that inter-

fered with or facilitated a subject’s use of the apparatus

during the subsequent trial. It was not feasible to clean the

apparatus (which were large, difficult to handle and store,

and included porous materials) between trials, and cleaning

agents are strictly regulated by the Safari Park. We think it

is unlikely that any odors that Swazi may have deposited

on the apparatus prior to or during a modeled trial would

have been particularly salient, given that the apparatus was

used repeatedly by the subjects, touched by keepers, and

stored on the ground in areas where they may have come in

contact with other olfactory stimuli. Nonetheless, we

reviewed the tapes for evidence that there may have been

olfactory biases in the data. For example, if Swazi’s scent

cues were salient for subjects in the modeled condition, we

should expect to see higher focus scores and initial interest

in subjects who received apparatus that were more fre-

quently touched by Swazi during modeled sessions. This

was not the case. Likewise, Swazi’s scent cues also might

have directed subjects toward Swazi’s chosen solution if

the majority of her scent cues were in proximity to parts of

the apparatus relevant to that particular solution. This also

appears not to be the case. As mentioned previously, when

demonstrating an apparatus, Swazi’s touches were always

Fig. 8 Initial interest is illustrated above in unmodeled and modeled

conditions

Anim Cogn (2013) 16:459–469 467

123

weighted (at least 2:1) toward the solution mechanism she

demonstrated, whereas the attention of the subjects was

more evenly distributed. Furthermore, the subjects failed to

solve the apparatus via Swazi’s chosen mechanism on nine

of the 13 trials. Thus, while we cannot rule out odor cues,

we do not believe that they accounted for the pattern of

results we obtained.

The failure of our elephants to demonstrate copying

fidelity may have been related to the nature of the exper-

imental task. Inspired by studies with primates, we used

apparatus that required the elephants to manipulate con-

tainers that held hidden food rewards. This presents a sit-

uation that is more ecologically valid for primate species

(e.g., nut cracking, seed extraction, and ‘‘fishing’’ for

insects) than it is for elephants, whose foods do not typi-

cally require processing (Johnston 1981; Tomasello and

Call 1994; Sukumar 2003; Hart et al. 2008; Hermes et al.

2008). Furthermore, adult elephants probably only rarely

come across novel foraging situations, limiting the need for

social transmission of new feeding techniques. Elephants

develop the majority of their feeding/foraging skills within

the first year of life (Sukumar 2003), suggesting that

immature elephants may be the best subjects to use when

looking for evidence of true imitative social learning in

feeding contexts. Also, we conformed with primate studies

of social learning by using vision as the sensory modality

in which we tested the elephants. Although we attempted to

account for the elephants’ rather poor visual acuity (Shyan-

Norwalt et al. 2010) by building large apparatus and sta-

tioning the observing subjects close to the model, it may be

the case that elephants rely primarily on their superior

olfactory and/or auditory senses to glean information in

social situations and are unused to visually scrutinizing

their groupmates’ behavior. Studies of social learning

should try to tap the most appropriate sensory modalities

for the species under investigation.

In sum, our data suggest that elephants may benefit from

social learning, but not necessarily in the form of imitating

the actions of conspecifics in foraging contexts. Adult ele-

phants may learn from one another primarily in non-imitative

ways such as observational conditioning. Ours is the first

study to attempt a controlled investigation of social learning

in elephants, and our data should be taken with some caution,

considering the small sample and variables that we could not

control, such as possible odor cues. In future research, it will

be important to consider the ecological validity of social

learning experiments with elephants.

Acknowledgments We are very grateful to the elephant training

staff at the San Diego Zoo Safari Park for their assistance, support,

and encouragement, and to Melissa Ritzer and Erin Lane for their

diligent coding of the videotapes. Dr. Doree Fragaszy generously

offered advice on an earlier version of this paper, as did two anon-

ymous reviewers.

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