1

Personalities and problem-solving abilities of black-capped chickadees

(Poecile atricapillus).

Penghui (Carrie) Sun

A thesis submitted to the Department of Biology in partial fulfilment of the requirements

for the requirements for the degree of Bachelor of Science (Honours)

Queens University
Kingston, Ontario, Canada
April 2014

ABSTRACT
The relationship between personalities and problem-solving abilities of individual blackcapped chickadees (Poecile atricapillus) was assessed experimentally using birds in
captivity. Personality, a set of distinct behavioural traits that distinguishes individuals, was
scored and ranked for each of three chickadees. With respect to personality traits, bird G
was most exploratory and moderately neophobic, bird W was moderately exploratory and
least neophobic, and bird B was least exploratory and most neophobic. Following
personality assessment, each of the subjects were given 30 minutes to solve a foil topsealed container containing a visible waxworm reward. They were given 2 types of the
device: one with an open bottom and one with a sealed bottom. Bird B was unsuccessful in
the 1st trial with an open bottom, but succeeded with an average of 543 s with all other trials
(trial 2 with open bottom, and trial 1 and 2 with sealed bottom). Bird G was successful on
all trials with an average solving time of 250 s, and bird W was unsuccessful with all trials,
except the 2nd trial with an open bottom, solving it in 372 s. A second problem was given to
the subjects, a tube with a push-lever that released a visible wax-worm reward, but none of
the birds were able to solve this within the 30 minutes allotted for each trial. Overall, G was
the most successful solving the problems, while W was the least successful. I conclude that
individual personalities of black-capped chickadees may have a complicated effect on their
ability to persist and solve some problems, with various combinations of certain personality
traits being more successful. Variation in personality characteristics may extend to learning
speed in black-capped chickadees. This study gives insight into developing behavioural
assays for future studies on avian personalities.

ACKNOWLEDGEMENTS
Thank you to the following individuals for their dedication, help, and teamwork in helping
care for the chickadees: Lora Walsh, Lydia Rocheleau.
Thank you to Laura Nagel for being a supportive committee member for this project, and
giving helpful, constructive feedback.
Special thanks to Marianne Gagnon for your dedication in helping care for the chickadees
and being a major help in performing trials.
Biggest thanks to the supervisor of this project, Dr. Robert Montgomerie, who provided this
amazing opportunity to get to work with him and birds. Thank you for being so patient,
accommodating, and supportive throughout this journey.

TABLE of CONTENTS
Title
List of Figures & Tables
Introduction and Literature Review
Personality in birds
Problem-solving in birds
Relationships between personality and problem-solving in birds
This study
Methods
Study animals
Assessing personality
Assessing problem-solving
Results
Personality
Problem-solving
Discussion
Personality
Problem-solving
Relationships between personality and problem-solving
Limitations
Future Directions
Literature Cited
Summary
Appendix

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LIST of FIGURES & TABLES

Figure or Table
Figure 1a. Novel environment A, for personality assessment of exploratory

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behaviour.
Figure 1b. Novel environment B, for personality assessment of neophobic

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behaviour
Figure 1c. Novel environment A in trial (with home cage attached)
Figure 2a. Transparent Plexiglas foil-top container, with visible waxworm

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inside, with open bottom.

Figure 2b. Transparent Plexiglas foil-top container, with visible waxworm

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inside, with sealed bottom.

Figure 2c. Transparent Plexiglas tube device, with pushable lever to drop down

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the visible waxworm on top of platform.

Figure 3. Exploratory trial Number of location changes in novel environment

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(cage top, sides, floor, perch 1, 2, 3, feeder).

Figure 4. Exploratory trial Proportion of time in home environment.

Figure 5. Neophobia trial Proportion of time spent at feeder with rabbit.
Figure 6. Neophobia trial Number of times visited feeder with rabbit.
Figure 7. Neophobia trial Number of times pecking rabbit.
Figure 8. Neophobia trial Proportion of time each bird spent at each feeder.
Figure 9. Solving times of each trial and bird for foil-top problem.
Table 1a. All behaviours noted in exploratory and Neophobia trials.
Table 1b. List of behaviours deemed as important in contributing to personality.
Table 2. Problem-solving ability of subjects (solving time if successful and
method used).

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INTRODUCTION & LITERATURE REVIEW

Individual birds have distinctive personalities, a set of distinct behavioural characteristics
that distinguishes each of them from other members of the population or species. To
characterize and quantify personality in a bird subject, researchers typically use
assessments of exploratory behaviour and neophobia (fear of novel objects or
environments). They then use analyses to score or rank the subjects behaviour in terms of
characteristics. For example, frequent visits to a novel feeder may indicate boldness and
being more exploratory. A birds personality will often be quantified as either being
proactive (bold, aggressive, fast-exploring) or reactive (shy, cautious, slow-exploring).

Personality in birds
Personality can influence or predict other factors in a bird. Alpin et al. (2013) performed an
assessment of personality of different individuals in a flock of great tits (Parus major), and
found that the birds personalities determined the social organization of the flock. Assays of
exploratory behaviour in a novel environment with 5 artificial trees were used to quantify
their personality as either proactive or reactive. The types of behavioural observation
included number of flights, flight duration, and substrates they landed on. Social phenotype
was then measured using individual network measures: degree centrality, betweenness

centrality, and average association strength. Degree centrality is the number of other
individuals with which an individual has been observed associating with. Betweenness
centrality is the number of shortest path vertices to all other individuals that pass through
the focal individual. Average association strength is the average proportion of foraging time
spent with each of its associates, divided by an individuals association strength by its
degree. Then, a social network was mapped with all the individuals of the flock, including
how pro-active and reactive they were. Pro-active individuals had more social
associates than reactive individuals, but reactive birds formed more stable
relationships. They also found that males tended to associate with birds of a similar
personality type, but females did not show the same association pattern. Thus, the birds
personalities influenced their flock social structure and organization. These differences in
personality were found to have a genetic basis, thus can also be acted on by selection. This
also provides implications for the strength and direction of selection on personality in
natural populations.
Personality also plays a large role in mate choice and sexual signaling. Partners
should develop preferences for personalities that maximize reproductive output. Bird song
has a prominent and well-established role in sexual selection. Thus, Garamszegi et al.
(2008) studied the relationship between individual birds songs and their exploratory and
risk-taking behaviours in a field study of wild male collared flycatchers (Ficedula
albicollis). To do this they characterised personality based on the exploration behaviour at a
modified nest box, and on flight distance taken when a potential predator was approaching
during a simulated territorial interaction. They quantified an individuals song by

characteristics including its length, rate, complexity, repertoire size, and post (position at
singing relative to the surrounding vegetation). After calculating correlations between song
characteristics and exploration/risk-taking behaviour, they found that explorative and risktaker individuals consistently sang at lower song posts than shy individuals in the presence
of a human observer. Males from lower posts established pair-bonds relatively faster than
males from higher posts. These results may demonstrate that personality may be manifested
in bird song and play a prominent role in sexual selection.
Personality can be related to a birds lifestyle and environment. Mettke-Hofmann et
al. (2002) demonstrated this in parrot species (Pssitacidae), and showed that differences in
exploratory behaviour and neophobia affects how they interact with their environment.
They predicted that personality should differ between species if it constitutes an adaptation
to different environmental conditions. Sixty-one parrot species from eight clades with
different diets and habitat preferences were investigated in aviaries. In the exploration test,
a novel object (wooden ring) in the familiar aviary was presented. Latencies until first
contact with the object and the duration of exploration were recorded. In the neophobia test,
novel objects were placed beside the feeding dish and latencies until first food intake were
recorded. They performed multiple regression analyses with the data and compiled
phylogenetic relationships. They found that species that inhabit complex habitats, or that
feed on buds or species from islands, showed the shortest latencies in the exploration test.
In contrast, long latencies were related to a diet including a great amount of seeds and/or
flowers. The longest duration of exploration occurred in species eating nuts or originating
from islands, whereas short durations were related to feeding on seeds. Neophobia was

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positively related to a diet consisting of insects, and negatively to a diet of leaves. Thus,
personality in parrots seem to be strongly related to their species-specific ecology.
Personality also can be related to a birds physiology. Cockrem (2007) found that
individual corticosterone and behaviour responses depend on each birds personality. He
looked at mean and individual plasma corticosterone responses to handling of Japanese
quail (Coturnix japonica), broiler chickens (Gallus gallus domestincus), Adelie penguins
(Pygoscelis adeliae), and great tits. Quail and broiler chickens were sampled at a
commercial farm. Adelie penguins were caught on nests on Ross Island, Antarctica. Wild
great tits were caught and held individually in cages for 10 days. To induce stress, birds
were removed from their cages, handled for 15 minutes, and then held individually in boxes
for another 45 min. Corticosterone secretion from the cortex of the adrenal gland in birds
increases one or two minutes after the hypothalamo-pituitary-adrenal axis is activated in
response to stressors. To measure the birds personalities, birds were released individually
into a novel room or presented with novel objects in their cage. The rates of exploration in
the room and approach to the novel object are scored and used to classify birds as proactive
or reactive explorers. Birds with proactive personalities had relatively low corticosterone
stress responses, whilst birds with reactive personalities had large corticosterone responses.
Relationships between the physiological and behavioural characteristics of avian
personalities can now be explored in detail to determine the significance of individual
differences in stress responses and personalities in birds.

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Problem-solving in birds
Personality may also influence cognitive traits in birds, such as problem-solving. Thus
several species of birds are able to solve tasks of varying degrees. One of the most famous
cases was the observational study of the blue tits (Poecile caeruleus) in England in the
1920s-40s, who learned to peck through foil tops of milk bottles to obtain the milk reward
inside (Fisher and Hinde 1949). The innovation was first noticed in 1921 in Swaythling, a
town near Southampton, England, and noticeably spread throughout the country. These
milk bottles were available outside many peoples porches, when the milkman delivered
them to homes. Fisher and Hinde (1952) have suggested that the actions used in opening
the bottles might have been derived from innate motor patterns. The movements used were
the same used when feeding on natural foods. For example, the pecking motion to break the
foil seals is similar to the motion in opening nuts, and the tearing motion often seen is
similar to the movement used in tearing bark from a twig. They also suggest that initial
discovery of the bottle as a source of food may have been due to their natural feeding
habits. Both blue and great tits inspect many objects that appear to contrast with their
surroundings and often test their palatability. If the object is unpalatable, they sometimes
try to open it by hammering, which they will continue trying if they hear the object is
hollow. Thus, by following this pattern of motions, it just so happened that milk bottles
yielded a reward. Then all the birds had apparently learned to do this by local enhancement,
a type of social learning where attention is directed towards a place/object that another
animal is interacting with, which improves their individual learning.

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The phenomenon of opening of foil top bottles has also been seen in several other
studies. In a controlled experimental setting with black-capped chickadees (Parus
atricapillus), Sherry and Galef (1984) found that chickadees that fed from previously
opened cream bottles, and chickadees that observed trained conspecifics open and drink
from cream bottles, were more likely to learn to open sealed cream bottles than were nave
chickadees. Direct observation of behaviour of conspecifics might facilitate learning by
nave chickadees. The authors proposed several reasons why this might happen, such as the
idea that presence of a trained conspecific may have reduced fear (Clayton 1978) or
vigilance (Elgar 1989).
The presence of conspecifics seems to have a marked effect on problem-solving.
Liker and Bokony (2009) tested this by presenting an opening of feeder problem with house
sparrows (Passer domesticus) in Hungary in small and large groups (2 versus 6 birds). They
found that group size had strong effects on problem solving: sparrows performed 4 times
more and 11 times faster openings in large than in small groups, and all members of large
groups profited by getting food sooner (7 times sooner, on average). They also found that
urban groups were more successful than rural groups. The authors suggest that larger
groups may contain more diverse individuals with different skills and experiences, which
may increase the chance of solving the task by some group members. Increased success in
problem solving may promote group living in animals and may help them to adapt quickly
to new situations in rapidly-changing environments.
Group learning has also been seen in mixed-species flocks of chickadees. Krebs
(1973) found that two species of chickadees, chestnut-backed (Poecile rufescens)

13

chickadees and black-capped chickadees, learn from one another about the location and
nature of potential feeding places when they are foraging together in mixed flocks. When
an individual finds a single food item or is unsuccessful in finding a food item, members of
the other species modify their foraging behaviour by putting more effort into searching near
the site of the find. This phenomenon was seen in both species. They also found that when
two species are trained to forage in different places in the experimental trees, they converge
in their foraging behavior when they are put in mixed flocks, due to copying. Moreover,
individuals of both species are more likely to discover a completely new foraging place if
they are in the presence of an experienced bird of the other species. Therefore, learning
about potential feeding places from other species is an important function of mixed flocks,
at least for some species. Thus, these variations in innovation and problem-solving skills
may be naturally selected for under certain conditions.
Sasvari (1979) found that observational learning is often dependent on success in
certain environments, thereby dependent on species, when he compared the ability of great
tits, blue tits, and marsh tits (Poecile palustris) to learn food finding from other individuals.
To do this, he used a device in which the bird had to lift a linen cloth to reach for a
mealworm inside. The great tits were more successful at learning from conspecifics than
the other two species. This seems to be related to difference in adaptability in the wild,
where the great tit is most successful in urban environments: the great tit is known to take
advantage of man-made food sources and nesting materials, while blue tits only nest in
parks and large gardens and marsh tits almost never nest in cities at all.

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Problem-solving can differ in various degrees of difficulty, in which some species

are more versatile and innovative in solving more difficult levels of problems. Ravens
(Corvus corax), keas (Nestor notabilis), and New Caledonian crows (Corvus moneduloides)
have been shown to successfully solve various difficult food tasks, such as string-pulling,
where birds must attempt to pull up a string attached to a perch to obtain a piece of food
hanging from the other end. Given meat on a string for the first time, five out of six oneyear-old ravens tried several ways to reach the food, and eventually showed a coherent
sequence of pulling up and stepping on loops of string, leading to success in solving the
problem. Another task was given to them in the same study in which they had to use a
pulley device by pulling one end of a string to lift the meat attached to the other end of the
string. The ravens who were successful in the first string-pulling task also succeeded in the
pulley task (Heinrich and Bugnyar 2005). Keas were also successful at completing the
string-pulling task. When given the problem, the seven kea subjects showed immediate
interest in the string and, with the exception of a fledgling, successfully solved it and
reached the food by repeating combined actions of pulling up the string with the beak and
holding loops of it against the perch with a foot (Werdenich and Huber 2004). New
Caledonian crows could solve the problem as well. Taylor et al. (2010) looked at the
cognition behind this problem-solving. They found that when visual feedback was available
via a mirror mounted next to the string-solving apparatus, untrained crows performed at the
same level as an experienced crow, previously trained with similar string-pulling tasks.
Innovative tool use has also been seen in birds. Bird and Emery (2009) have shown
that rooks, which do not appear to use tools in the wild, can innovatively bend a piece of

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wire into a hook and use it to obtain an otherwise unreachable food reward. Furthermore,
dominant rooks (Corvus frugilegus) solved tasks faster than subordinate rooks. New
Caledonian crows were also able not only use sticks to their advantage when presented with
them, but also solve a sequenced problem requiring multiple steps (Taylor et al. 2010).
They could obtain a short stick by pulling a string, use the short stick to take out a long
stick from a toolbox, and finally use the long stick to take out food from a hole. Thus, New
Caledonian crows can solve complex physical problems by reasoning both causally and
analogically about causal relations, even when it concerns the functional properties of novel
types of tool. It is also evident that New Caledonian crows understand the abstract idea of
out of reach objects can be accessed using a tool. Causal and analogical reasoning may
form the basis of the New Caledonian crows exceptional tool skills (Taylor et al. 2009;
Taylor et al. 2010a; Taylor et al. 2010b).

Relationships between personality and problem-solving

There has been limited research regarding the link between personality and problemsolving. For example, Cole et al. (2011) assessed personality and individual variation in
problem-solving performance in great tits. They used neophobia trials to assay the subjects
personalities, by measuring their latency in approaching 2 feeders, one with a novel object
and one without, and taking a food from one of the feeders. A pull-out lever problem was
then given to each of the subjects to solve. To obtain a visible food reward, the bird had to
pull out a lever that would cause the food to drop down. It was found that neophobia did

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not have a correlation with how well the birds performed in the task. This suggests that
individual differences in problem-solving may represent an independent source of
behavioural variation in natural populations, at least for great tits.
However, Cole et al.s (2011) study seems to have results contrary to those from
most other studies that seem to suggest that personality has an influence on problemsolving ability. After assessing personality in 3 passeriforme species, the Carib grackle
(Quiscalus lugubris), the lesser Antlillean bullfinch, (Loxigilla noctis), and the shiny
cowbird (Molothrus bonariensis), and 2 columbiformes, the Zenaida dove (Zenaida aurita)
and the common ground dove (Columbina passerina), neophobia was found to be
influential in determining their response and ability in solving a feeding problem. Given a
Plexiglas box to retrieve a visible food reward, less neophobic birds were faster at
successfully solving the box than more neophobic birds. Also, passeriformes were faster at
solving than columbiformes (Webster and Lefebvre 2000). Similar results were obtained
with black-capped chickadees. Roth et al. (2010) examined the link between personality
and problem-solving ability of 2 populations of black-capped chickadees, one population
from Alaska (a harsher environment) and one population from Kansas (a milder
environment). They assessed personality by neophobia trials, measuring each subjects
latency in approaching a novel feeder and taking a food from it. They gave each subject a
problem in which the bird was to push a weighted washer off a well to retrieve a visible
food reward. It was found that Alaskan birds were both less neophobic and faster at solving
the well task. These results are consistent with another black-capped chickadee study,
which it was determined that chickadees that engage in more exploratory behaviours in a

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novel environment learned an acoustic discrimination task faster than birds less exploratory
in a novel environment. The novel environment used was a room consisting of 5 artificial
trees. A trend was seen in which birds that visited more trees during their exploratory trial
took fewer problem-solving trials to learn the acoustic discrimination task (Guillette et al.
2009).
There may be other factors that contribute to personality, problem-solving, or both.
For example, the bigger forebrain size in many species of North American and British Isles
birds is positively correlated with better foraging innovations with food problems (Lefebvre
et al. 1995). Two data sets were collected on opportunistic foraging innovations of birds in
North American (1973-1993; N=196 species) and the British Isles (1983-1993; N=126
species). They looked at foraging innovations for all orders of birds, and for both the
baseline of the forebrain and the brainstem of forebrain. Both body parts showed a positive
relationship between size and innovation rate. Trends between brain size and rate of
structural evolution are confirmed and suggest that innovation rate in the field may be a
useful measure of behavioural plasticity.

This Study
My study was designed to assess the relationship between personality of black-capped
chickadees and their ability to solve two simple tasks. Much of the literature supports the
idea that personality is influential in predicting problem-solving ability, with the exception
of Cole et al. (2011). Therefore for this study, I expected that more exploratory and less

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neophobic chickadees will be faster and more innovative at solving the tasks than less
exploratory and more neophobic chickadees.

MATERIALS AND METHODS

Study Animals
Three black-capped chickadees from the same population, and probably the same social
group, were captured using potter traps at Queens University Biological Station (Elgin,
Ontario) in September 2013. Each bird was banded a colour band, and named B, G, and W,
the colours of their bands. They were brought to and housed in the Animal Care Facilities at
Queens University (Kingston, ON). All three individuals were placed into their own
individual cages, with all three cages were kept in a temperature-controlled room with a
stable room temperature of about 20oC and dimensions of 3.66 m x 3.66 m. The room was
also light-controlled and was set to reflect the chickadees natural light-dark cycle. Each
cage was provided with 3 perches, a cuttlebone, and a UV lamp. Their cages were prepared
with vegetation clipped around and inside the sides. Each cage was of dimensions 0.45m x
0.45m x 1m. Cages were cleaned and replaced once a week. Chickadees were fed once each
morning. Their food was mixed using a coffee blender, in a ratio of 1 part of sunflower
seeds, 1 part Mazuri small bird seed, and 2 part peanuts. One scoop of bird seed was
placed over top the mixture. Birds were fed 1 tablespoon of this mixture daily. Birds were
also supplied with 3 clean bowls of water to bathe and drink every morning.

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Assessing Personality
To assess personality, I set up 2 novel environments in a separate trial room. The trial room
was kept at a controlled temperature of about 20oC, and was 2.74 m x 2.44 m. Novel
environment A was for assessing exploratory behaviour and was an empty cage with 3
tiered perches, newspaper covering the bottom, 1 familiar food feeder with 1 scoop of bird
seed and 1 water feeder half-filled with water (Figure 1a). Novel environment B was more
specifically for assessing neophobia of objects, and consisted of 3 tiered perches,
newspaper covering the bottom, 1 water feeder half-filled with water, 1 familiar food feeder
with 1 scoop of bird seed, and 1 food feeder with 1 scoop of bird seed and had a small
stuffed rabbit attached to the cages side, hovering above the feeder (Figure 1b). Each
subject was individually transferred to novel environment A the first day, and then to the
novel environment B the second day. Before being transferred to either novel environment,
birds were starved (i.e. the food dish from their home cage was removed) 1 hour before. To
transfer them, a black blanket covering the novel environment was switched onto their
home cage to entice them to go toward the light and into the novel environment. Each day
had a randomized order of subjects being transferred. Each subject was given 30 minutes
with the novel environment they were transferred to, but the subjects home cages remained
attached throughout the 30 minute trial, allowing for flying back and forth and actions in
either cage (Figure 1c). All trials were videotaped.

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Figure 1a. Novel environment A

Figure 1b. Novel environment B

Figure 1c. Novel environment A in trial (with home cage attached)

The videos were watched and behaviours were timed using the JWatcher program
(v. 0.9; Los Angeles, USA and Sydney, Australia). During assessment, I watched the videos
and called out the behaviours as they happened, while an assistant pressed the keyboard
entries corresponding to the behaviours. Note that there may be slight human error with

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regards to scoring behaviours using JWatcher, as sometimes the birds move faster than the
human eye can catch their behaviours. The program logged all data, including but not
limited to: time taken between behaviours, total number of times a behaviour happened,
average number of times a behaviour happened, average length of time performing a
behaviour, total time performing a behaviour, and proportion of time taken performing a
behaviour.
Table 1a. All behaviours noted in exploratory and Neophobia trials
Exploratory trials
Cage back
Cage front
Cage right
Cage top
Feeder
Feeding
Home floor Home perch
Home water Perch 1
Perch 3
Preening
Chickadee-call

Cage left
Drinking
Floor
Home side
Perch 2
Tsit call

Neophobia trials
Rabbitfeeder Peck rabbit
Cage front
Cage left
Cage top
Drinking
Feeding
Floor
Home perch Home side
Perch 1
Perch 2
Preening
Tsit-call

Cage back
Cage right
Feeder
Home floor
Home water
Perch 3
Chickadee-call

Using JMP (v. 11.0.0; Cary, North Carolina), ANOVA analyses followed by a
Tukey-Kramer comparison were performed to determine if significant rends for a relevant
behaviour. If so, that behaviour was counted as an important variable that contributed to the
birds personalities. A behaviour was included in my personality assessment if both other
thesis student Lora Walsh (B.Sc. [Hons.]) and I agreed that (i) the trend of the behaviour
contributed to our understanding of its personality and (ii) it showed a consistent trend for a
particular bird. For example, number of times pecking the rabbit was selected because it
suggested less neophobia within a bird and the same pattern was seen for each bird across

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my and Walshs trials (i.e. Bird G was highest for both trials). These significantly relevant
behaviours were evaluated later in the context of their problem-solving results.

Assessing Problem-Solving
The first problem-solving task consisted of giving each bird two kinds of clear Plexiglas
containers taped with foil, containing a visible waxworm reward. For the first trial, birds
were given the device with an open bottom (Figure 2a), giving the bird two ways to solve
it, either knocking it over or pecking through the foil. The second kind had a sealed bottom
(Figure 2b), forcing the bird to only solve it by pecking through the foil. For each trial,
birds were given a maximum of 30 minutes to solve it. Each of these 2 kinds of trials were
replicated. The order of which subjects were given the task was randomized.
4 cm
Aluminum foil taped on top

2 cm
Waxworm reward

Figure 2a. Transparent Plexiglas foil-top container, with visible waxworm inside, with
open bottom.
4 cm
Aluminum foil taped on top

2 cm

Paper taped on bottom

Waxworm reward

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Figure 2b. Transparent Plexiglas foil-top container, with visible waxworm inside, with
sealed bottom.
Lever to push

4 cm
Waxworm reward

The second problem-solving task

consisted of giving each bird a clear
Plexiglas tube containing a lever
mechanism that when pushed, turns a
platform holding a visible waxworm
reward, making the waxworm fall below
on a ledge where the bird can eat it
(Figure 2c). Each bird was individually
transferred to a cage containing this
device, and given 30 minutes to attempt

this problem once.

18
cm

Figure 2c. Transparent Plexiglas tube device, with pushable lever to drop down the visible
waxworm on top of platform.

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RESULTS
Personality
Many behaviours were assessed (Table 1a) in the trials, but only a few behaviours were
deemed relevant and applicable to assessing exploratory behaviour and neophobia (Table
1b). See METHODS pages 21-22 for why and how behaviours were chosen.
Table 1b. List of behaviours deemed as relevant in contributing to personality
Exploratory trial behaviours
Proportion of time in home environment
Proportion of time spent exploring in the novel

High/More
Less exploratory
More exploratory

Low/Less
More exploratory
Less exploratory

environment (cage top, sides, floor, perch 1, 2, 3, feeder)

Neophobia trial behaviours
Proportion of time spent at rabbit feeder
Number of times visiting rabbit feeder
Number of times pecking rabbit
Proportion of time spent at familiar feeder

Less neophobic
Less neophobic
Less neophobic
More neophobic

More neophobic
More neophobic
More neophobic
Less neophobic

Exploratory trials:
Bird G and bird W were inconsistent between trials 1 and 2 for number of location
changes. They were almost equal in trial 1, but in trial 2, bird G had noticeably more
location changes. Most importantly, bird B consistently had the least number of location
changes between trial 1 and 2, compared to bird G and bird W (Figure 3). All birds were
relatively consistent between trials for proportion of time spent at home, in which bird G
spent the least proportion of time in the home environment, bird B spent the most time in

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the home environment, and bird W being moderately in the middle (Figure 4). See
APPENDIX (Figure A, B) for statistical results.

Figure 3. Exploratory trial Number of location changes in novel environment (cage top,
sides, floor, perch 1, 2, 3, feeder). Mean diamonds demonstrate a sample mean and a 95%
confidence interval. Sample size: N=3. (Note that the y-axis is plotted on a log scale).

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Figure 4. Exploratory trial Proportion of time spent in home environment. Mean

diamonds demonstrate a sample mean and a 95% confidence interval. Sample size: N=3.
Neophobia trials:
Bird B spent a slightly higher proportion of time than bird G at the rabbit feeder in
trial 1, but this proportion was almost equal in trial 2. Importantly, bird W spent a
significantly higher proportion of its time at the feeder with the rabbit, compared to bird B
and bird G, especially in trial 1. In trial 2, bird W still spent a higher proportion of time at
the rabbit feeder compared to bird B and bird G, but noticeably less compared to its
proportion in trial 1 (Figure 5). The trend of number of visits to the rabbit feeder matched
with the trend of proportion of time spent at the rabbit feeder. Bird B visited the rabbit
feeder slightly more times than bird G in trial 1, but this difference became negligible in
trial 2. For both trials, bird W also visited the feeder with the rabbit significantly more

27

times than bird B and bird G, but had visited slightly less times in trial 2 (Figure 6).
However, even though bird W visited the rabbit feeder the most, for both trials 1 and 2, bird
G consistently pecked the rabbit significantly more times than bird B and bird W. Bird W
spent pecked the rabbit slightly more times than bird B in trial 1, but this difference became
negligible in trial 2 (Figure 7). For both trials, bird W spent the highest proportion of time
interacting with both feeders, compared to bird B and bird G. Also for both trials, bird W
interacted more with the feeder with the rabbit compared to the familiar feeder. The
proportion of time bird W spent at both feeders was drastically higher in trial 1 compared to
trial 2. There were inconsistent results between bird B and bird G, and between their trials.
In trial 1, bird B spent a higher proportion of time at the feeder with the rabbit compared to
the familiar feeder, but for trial 2, spent no time at all there and spent more time at the
familiar feeder. In trial 1, bird G spent almost equal proportions of time split between the 2
feeders, but in trial 2, spent a higher proportion of time at the familiar feeder compared to
the feeder with the rabbit. Bird B spent a slightly higher proportion of time for both feeders
than bird G, except for trial 2 at the familiar feeder, when bird B spent no time at all there
(Figure 8). See APPENDIX (Figure D, E, F) for statistical results.

28

Figure 5. Neophobia trial Proportion of time spent at feeder with rabbit. Mean diamonds
demonstrate a sample mean and a 95% confidence interval. Sample size: N=3.

Figure 6. Neophobia trial Number of times visiting feeder with rabbit. Mean diamonds
demonstrate a sample mean and a 95% confidence interval. Sample size: N=3.

29

Figure 7. Neophobia trial Number of times pecking rabbit. Mean diamonds demonstrate
a sample mean and a 95% confidence interval. Sample size: N = 3.

Figure 8. Neophobia trial Proportion of time each bird spent at each feeder. Mean
diamonds demonstrate a sample mean and a 95% confidence interval. Sample size: N = 3.

30

Problem-solving
Each of the subjects were given 30 minutes to solve a foil top-sealed container
containing a visible waxworm reward. They did this once for a container without a bottom
and once for a container with a bottom. Bird B was unsuccessful in the 1st trial without the
bottom, but successfully used the knock-over method for the 2nd trial without the bottom,
and the pecking method for both trials with the bottom. These 3 successful trials had an
average of 543s solving time. Bird G was successful on all trials with an average solving
time of 250s. It used the knock-over method in the 1st trial without the bottom, and the
pecking method in all other trials. Bird W was unsuccessful with all trials, except the 2nd
trial without the bottom, solving it in 372s, using the pecking method. Bird B and bird G
showed signs of using multiple solutions to a problem, either pecking through the foil or
knocking it over if it had an open bottom. Bird W could not attempt different methods to
solving the problem and consistently tried the same strategy; it kept pecking at the sides.
Table 2 summarizes all successful solving times and what method the bird used to solve the
problem. Figure 9 shows the trend of each birds successful solving time for the problem.

31

Trial/Subject

Blue

Green

White

Trial 1, device with

Unsuccessful

2m05s (125s)

Unsuccessful

open bottom

knock-over

Trial 1, device with

17m07s (1027s)

11m15s (675s)

Unsuccessful

sealed bottom

pecking

pecking

Trial 2, device with

1m52s (112s)

1m50s (110s)

6m12s (372s)

open bottom

knock-over

pecking

pecking

Trial 2, device with

8m10s (490s)

1m30s (90s)

Unsuccessful

sealed bottom

pecking

pecking

Average solving time

543s

250s

372s

Table 2. Problem-solving ability of subjects (solving time if successful and method used)

Figure 9. Solving times of each trial and bird for foil-top problem. Sample size: N = 3.

32

Another problem was given to the subjects, a clear tube with a push-lever that
released a visible wax-worm reward, but none of the birds were able to solve this within the
30 minute allotted time span. All birds pecked at the tube and did not try manipulating the
lever in any such way.

DISCUSSION
Personality
Bird B had the least number of exploratory location changes in both trials (Figure 3),
demonstrating that bird B is not as exploratory as the other 2 birds in this aspect. For
proportion of time spent at home, all birds showed a consistent trend between trials, in
which bird G spent the least proportion of time in the home, bird B spent the most time in
the home environment, and bird W being moderately in the middle (Figure 4). Because
spending time in the home environment is equivalent to not exploring the new environment,
bird B again shows itself as least exploratory among the 3 birds, and bird G shows itself to
be relatively more exploratory than the other 2 birds. Thus, interpreting the data from
figures 3 and 4 altogether, I conclude bird G was most exploratory, followed by bird W,

33

then bird B (Figures 3, 4). Bird W spent a significantly higher proportion of time at the
rabbit feeder compared to bird B and bird G (Figure 5), suggesting bird W is least
neophobic. Furthermore, bird W spent a higher proportion of time at the rabbit feeder for
trial 1 over trial 2 (Figure 5). This suggests that bird W has lost a bit of interest in the novel
object since the first time it saw it in trial 1. Even so, bird W still spent a higher proportion
of time at the rabbit feeder for trial 2 than the other 2 birds (Figure 5), thus it still
demonstrates itself as consistently less neophobic than the others. Both bird B and bird G
spent a slightly lower proportion of time at the rabbit feeder in trial 2 compared to trial 1
(Figure 5), again likely because the birds lose interest in the novel object. Bird B spent a
slightly higher proportion of time than bird G at the rabbit feeder in trial 1, but this
proportion was almost equal in trial 2 (Figure 5). All these trends seen with number of visits
to the rabbit feeder matched with all trends of proportion of time spent at the rabbit feeder
(Figure 6). These trends seen in Figure 5 and 6 may suggest that bird B is less neophobic
than bird G, but results from all other neophobia trials suggest otherwise. Bird G
consistently pecked the rabbit significantly more times than bird B and bird W (Figure 7),
showing that bird G is not afraid of the novel object at all, and is bold enough to interact
with it. This suggests that bird G is least neophobic of all birds. Bird W spent pecked the
rabbit slightly more times than bird B in trial 1, but this difference became negligible in
trial 2 (Figure 7). The proportion of time bird W spent at both feeders was drastically higher
in trial 1 compared to trial 2 (Figure 8). Therefore, bird W seems to lose interest to novel
stimuli quickly (Figure 5, 7, 8) which may be important for the results of its problemsolving trials (see problem-solving section below). For both trials, bird W spent the highest
proportion of time at both feeders, compared to bird G and bird B, again demonstrating that

34

bird W is least neophobic. There were inconsistent results between bird B and bird G, and
between their trials. In trial 1, bird B spent a higher proportion of time at the feeder with the
rabbit compared to the familiar feeder, but for trial 2, spent no time at all there and spent
more time at the familiar feeder. In trial 1, bird G spent almost equal proportions of time
split between the 2 feeders, but in trial 2, spent a higher proportion of time at the familiar
feeder compared to the feeder with the rabbit. bird B spent a slightly higher proportion of
time for both feeders than Bird G, except for trial 2 at the familiar feeder, when bird B
spent no time there at all (Figure 8). It is unsure what contributed to these inconsistent
results for bird B and bird G, but I propose it demonstrates bird B and bird G are more alike
to each other than bird W, and are more unpredictable than bird W.
Thus, interpreting the data from figures 5-8 altogether, I conclude that bird W is
least neophobic, bird B is most neophobic, and bird G is moderately neophobic (Figures 58).
Overall, I have interpreted each birds personality based on 2 facets: how
exploratory they were in a novel environment and how neophobic they were in the presence
of a novel object. I conclude that: Bird B is least exploratory and most neophobic, bird G is
most exploratory and moderately neophobic, and bird W is moderately exploratory and
least neophobic.

Problem solving

35

It was then determined that bird G was most successful in solving the foil top problem, with
an average of 250s. Bird G shows a slight trend in learning as well, with its solving times
decreasing over trials overall. Bird B was next most successful, solving 3 out of 4 of the
trials with an average solving time of 543s. Bird W was least successful, being able to solve
only 1 out of 4 trials, solving it in 372s. Therefore, I confirm that black-capped chickadees
are able to solve foil-top tasks and able to show some degree of learning. Bird B and bird G
showed signs of using multiple solutions to a problem, either pecking through the foil or
knocking it over if it had an open bottom. Bird W could not attempt different methods to
solving the problem and consistently tried the same strategy. I attribute this to its
personality it seems to lose interest in stimuli and the task at-hand very quickly (Figure 5,
7, 8). This discrepancy demonstrates the variation that individual birds of the same species,
have in their innovation, learning, and problem-solving abilities.
None of the birds were able to solve the lever-pushing tube task, and none of the
birds showed any signs that they were making progress. The difficulty level of the leverpushing tube task proved to be too high for them. Thus, I conclude that black-capped
chickadees are able to solve problems only up to a certain degree of complexity.

Relationships between personality and problem-solving

Each of the birds had distinct personalities and each of the birds were able to solve
the food problems with varying degrees of success. This further demonstrates the idea that
a birds personality can impact other parameters of their lifestyle. I found that a very

36

exploratory and moderately neophobic personality seems to contribute to the best problemsolving ability in black-capped chickadees. A different combination of traits, a moderately
exploratory and hardly neophobic personality, seems to contribute to worst problem-solving
ability. Furthermore, if a bird loses interest in novel stimuli and tasks quickly, this seems to
contribute to its poor success, like bird W. It also seems that more predictable birds (bird B
and G, compared to bird W) are more successful at problem-solving and using various
innovations. This combination of traits that links to the best problem-solving ability is
different from most previous literature results, in which most exploratory and least
neophobic birds were best at problem-solving (Webster and Lefebvre 2000; Guillette et al.
2009; Roth et al. 2010). Therefore, the conclusions drawn from this study may suggest 2
ideas. The first is that individual variation in problem-solving is independent of differences
in neophobia and exploratory behaviour, a similar result to the study performed by Cole et
al. (2013). This then suggests that individual differences in problem-solving may represent
an independent source of behavioural variation in the natural population of Ontario blackcapped chickadees. The second idea these results could suggest is that the relationship
between personality and problem-solving may be more intrinsic and less clear as previous
literature showed. There may be a specific combination of traits in a personality that give
the most ideal problem-solving ability for example, being both very exploratory, but
slightly neophobic. Because natural selection acts on variation, this would imply evolution
may select for certain personality traits or combinations of traits, because these traits confer
fitness value in cognitive ability or adaptive techniques for food foraging. The results
suggest that variation in personality characteristics may extend to learning speed in blackcapped chickadees. These results would also suggest that there may be linkage with some

37

related genes, whether they be between different personality traits or between certain traits
and problem-solving. However, much more research needs to be done for these ideas to be
conclusive.

Limitations
This study has a few limitations that may prevent the results and interpretation of the results
from being completely conclusive. The biggest one is perhaps the limited sample size of 3
birds. Due to limited resources, the lab was only able to capture and maintain 3 birds.
However, an advantage of this is that I was able to perform the study comprehensively and
in-depth for the 3 individuals. Another limitation is that all data and results are subjective
and open to interpretation. This is a common limitation with many ethology studies, but
must be acknowledged regardless.

Future Directions
It is recommended that this study be replicated, but with a larger sample size. This can help
see if the results and the data are robust and perhaps make the studys results more
conclusive. With a larger sample size, more analyses and statistics can be done as well, and
a principal component analysis becomes a possibility as well. It may also be worthwhile
comparing personalities and cognitive abilities between 2 or more populations of the same
species, as Roth et al. (2010) did. This could provide more insight and a clearer picture of

38

the individual differences in birds. When results become more conclusive, it would be
worthwhile to at the genetic components of personality traits and problem-solving. Sex
differences may also be worthwhile to assess.
Although the study has its limitations and may not have provided completely
conclusive results, it has furthered the literature. The study has provided behavioural assays
for relevant studies in the future and given insight into selection of personality and
problem-solving traits, suggesting a complex relationship between the 2 factors. With
further research, literature can continue to be added and applications such as improving
breeding programs for chickadees and related birds can be implemented.
LITERATURE CITED
Alpin, L.M., Farine, D.R., Morand-Ferron, J., Cole, E.F., Cockburn, A. and Sheldon, B.C.
2013. Individual personalities predict social behaviour in wild networks of great tits
(Parus major). Ecology Letters, 16(11): 1365-1372.
Bird, C.D. and Emery, N.J. 2009. Insightful problem-solving and creative tool modification
by captive nontool-using rooks. Proceedings of the National Academy of Sciences,
106(25): 10370-10375.
Clayton, D.A. 1978. Socially facilitated behaviour. The Quarterly Review of Biology, 53:
373-392.
Cockrem, J.F. 2007. Stress, corticosterone responses and avian personalities. Journal of
Ornithology, 148: 169-178.

39

Cole, E.F., Dominic, L.C. and Quinn, J.L. 2011. Individual variation in spontaneous
problem-solving performance among wild great tits. Animal Behaviour, 81: 491498.
Dan Blumsteins lab and The Animal Behaviour Lab. 2012. JWatcher: v. 0.9. University of
California, Los Angeles and Macquarie University, Sydney.
Elgar, M.A. 1989. Predator vigilance and group size in mammals and birds: a critical
review of the empirical evidence. Biology Reviews, 64: 13-33.
Fisher, J. and Hinde, R.A. 1949. The opening of milk bottles by birds. British Birds, 42:
347-357.
Fisher, J. and Hinde, R.A. 1952. The opening of milk bottles by birds. Letters to Nature,
169: 1006-1007.
Garamszegi, L.Z., Eens, M. and Torok, J. 2008. Birds reveal their personality when singing.
PLoS ONE, 3(7): e2647.
Guillette, L.M., Reddon, A.R., Hurd, P.L. and Sturdy, C.B. 2009. Exploration of a novel
space is associated with individual differences in learning speed in black-capped
chickadees, Poecile atricapillus. Behavioural Processes, 82: 265-270.
Heinrich, B. and Bugnyar, T. 2005. Testing problem-solving in ravens: String-pulling to
reach food. Ethology, 111: 962-976.
Krebs, J. R. 1973. Social learning and the significance of mixed-species flocks of
chickadees (Parus spp.). Canadian Journal of Zoology, 51: 1275-1288.

40

Lefebvre, L., Whittle, P., Lascaris, E. and Finkelstein, A. 1995. Feeding innovations and
forebrain size in birds. Animal Behaviour, 53: 549-560.
Liker, A. and Bokony, V. 2009. Larger groups are more successful in innovative problem
solving in house sparrows. Proceedings of the National Academy of Sciences,
106(19): 7893-7898.
Mettke-Hofmann, C., Winkler, H. and Leisler, B. 2002. The significance of ecological
factors for exploration and neophobia in parrots. Ethology, 108: 249-272.
Roth, T.C., LaDage, L.D. and Pravosudov, V.V. 2010. Learning capabilities enhanced in
harsh environments: a common garden approach. Proceedings of the Royal Society
B, 277: 3187-3193.
SAS Institute Inc. 2013. JMP Statistical Software: v. 11.0.0. Cary, North Carolina.
Sasvari, L. 1979. Observational learning in great, blue, and marsh tits. Animal Behaviour,
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Sherry, D.F. and Galef, B.G. 1984. Cultural transmission without imitation: milk bottle
opening by birds. Animal Behaviour, 32: 937-938.
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41

Taylor, A. H., Hunt, G. R., Medina, F. S. and Gray, R. D. 2009. Do New Caledonian crows
solve physical problems through causal reasoning? Proceedings of the Royal
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2010. An investigation into the cognition behind spontaneous string pulling in New
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42

SUMMARY
1) The relationship between personality and problem-solving was assessed in a natural
population of Ontario black-capped chickadees.
2) Personality was assessed in the subjects using trials of exploratory behaviour and
neophobia. It was determined that Bird B was least exploratory and most neophobic,
Bird G was most exploratory and moderately neophobic, and Bird W was
moderately exploratory and most neophobic.
3) It was then determined that Bird G was most successful in solving and learning the
foil top problem, with an average of 250s. Bird B was next most successful, solving
3 out of 4 of the trials. Bird W was least successful, being able to solve only 1 out of
4 trials.
4) Thus, a very exploratory and moderately neophobic personality seems to be linked
to the best problem-solving ability in black-capped chickadees. A different

43

combination of traits, a moderately exploratory and hardly neophobic personality,

seems to be linked to the worst problem-solving ability.
5) The results may demonstrate one of two possible explanations. The first is that
individual variation in problem-solving may be independent of differences in
neophobia and exploratory behaviour. The second is that the relationship between
personality and problem-solving may be complex and intrinsic. There may be a
specific combination of traits in a personality that give the most ideal problemsolving ability. Because natural selection acts on variation, evolution may select for
certain personality traits or combinations of traits.
6) This study has limitations that may prevent the results and interpretation of the
results from being completely conclusive, such as the limited sample size of 3 birds,
and that all data and results are subjective and open to interpretation.
7) The study has provided behavioural assays for relevant studies in the future and
given insight into selection of personality and problem-solving traits, suggesting a
complex relationship between the 2 factors. With further research, literature can
continue to be added and applications such as improving breeding programs for
chickadees and related birds can be implemented.

44

APPENDIX
Statistics summaries for exploratory trials
A: Statistics for Figure 4 Number of location changes

45

B: Statistics for Figure 5 Proportion of time spent in home environment.

46

Statistics summaries for neophobia trials

47

C: Statistics for Figure 6 Proportion of time spent at rabbit feeder.

48

49

D: Statistics for Figure 7 Number of times visiting the feeder with the rabbit.

50

51

E: Statistics for Figure 8 Number of times pecking the rabbit.

52