Primates
DOI 10.1007/s10329-013-0360-8
NEWS AND PERSPECTIVES
Are shy individuals less behaviorally variable? Insights
from a captive population of mouse lemurs
Jennifer L. Verdolin • John Harper
Received: 20 December 2012 / Accepted: 10 May 2013
Ó Japan Monkey Centre and Springer Japan 2013
Abstract Increasingly, individual variation in personality
has become a focus of behavioral research in animal sys-
tems. Boldness and shyness, often quantified as the ten-
dency to explore novel situations, are seen as personality
traits important to the fitness landscape of individuals. Here
we tested for individual differences within and across
contexts in behavioral responses of captive mouse lemurs
(Microcebus murinus) to novel objects, novel foods, and
handling. We report consistent differences in behavioral
responses for objects and handling. We also found that the
responses to handling and novel objects were correlated
and repeatable. Lastly, we show that shyer individuals may
show less variability in their behavioral responses. This
study provides new information on the potential for
behavioral syndromes in this species and highlights
differences in the degree to which behavioral types (e.g.,
shy/bold) vary in their behavioral responses.
Keywords Boldness
Novel object
Personality

Behavioral repeatability
Temperament
J. L. Verdolin and J. Harper are co-first authors.
J. L. Verdolin (&)
National Evolutionary Synthesis Center (NESCent),
2024 W. Main Street, Durham, NC 27705, USA
e-mail: verdolin@nescent.org
J. Harper
Department of Biological Sciences, North Carolina State
University, Raleigh, NC 27695, USA
J. Harper
Department of Biological Sciences, University of Leeds,
Leeds LS2 9JT, UK
Introduction
Individual variation is the ‘grist’ of natural selection and
has implications for social dynamics, population structure,
and disease transmission (Drea and Wallen 1999; Réale
et al. 2000; Dingemanse and Réale 2005; Wilson 2007;
Hamede et al. 2009; Pinter-Wollman 2012). Despite this,
behavioral ecologists have traditionally considered indi-
vidual variation along a behavioral axis as statistical
noise. Recently, however, empirical evidence from a
broad range of taxa, including invertebrates and verte-
brates, has revealed that individuals may show consistent
differences that are correlated across different contexts
(Gosling 2001; Sih et al. 2004a, b; Bell 2005; Quinn and
Cresswell 2005; Powell and Gartner 2011), often referred
to as behavioral syndromes or personality (Gosling and
John 1999; Gosling 2001; Sih et al. 2004a, b; Wilson and
Godin 2009). As a consequence of fitness differences,
selection can and will sustain individual variation in a
population (Wilson 1998; Sih et al. 2004a), resulting in
polymorphic populations differing in life history traits
(Biro and Stamps 2008). Alternatively, differing life his-
tory strategies can produce variation in risk-taking
behavior among individuals within a population, thereby
promoting the evolution of personality (Wolf et al. 2007;
Dammhahn 2012).
Generally, individuals that approach novel objects and
food sooner, reflecting a willingness to explore, are con-
sidered bolder relative to others (Wilson et al. 1994; Bell
and Stamps 2004; Sih et al. 2004a, b). Although aggression
has also been correlated with boldness in a variety of
species (Wilson et al. 1994; Bell and Stamps 2004; Sih
et al. 2004a, b; Moretz et al. 2007; Pintor et al. 2008), it is
typically measured as conspecific aggression (but see Réale
et al. 2000). In contrast, the relationship between
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aggression and boldness, in the context of animal handling
by humans, may be more variable depending on the spe-
cies. For example, response to human handling may indi-
cate a fear response rather than conspecific aggression,
while a lack of response may indicate greater risk-taking,
or boldness (e.g., Möeller and Garamszegi 2012). There-
fore, responses to handling, or approach by humans,
reflects one important potential measure contributing to
behavioral syndromes (Réale et al. 2000; Möeller and
Nielson 2010; Möeller and Garamszegi 2012).
We investigated personality and variability in a captive
population of gray mouse lemurs (Microcebus murinus).
Gray mouse lemurs are small, arboreal, nocturnal pri-
mates found in the western and southwestern regions of
Madagascar (Martin 1972; Mittermeier et al. 1994;
Radespiel 2000). Strepsirrhine primates, such as the gray
mouse lemur, represent a primitive group and evidence of
behavioral syndromes in this group may suggest con-
vergence on this trait with other basal primate groups.
Recent experimental tests of wild populations demon-
strate consistent interindividual differences that were
correlated across exploratory behavior, foraging behavior,
and across differing fitness potentials (Dammhahn 2012;
Dammhahn and Almeling 2012). Here, we explored
whether captive gray mouse lemurs showed individual
variation in behavioral responses towards novel objects,
novel foods, and human handling. We then determined if
behavioral syndromes were present by assessing whether
responses were correlated across these contexts. First, we
predicted that we would detect individual variability.
Second, if behavioral syndromes exist in this species
using the behavioral axes we quantified, we predicted that
responses would be consistent within individuals across
contexts. Last, we predicted that there would be a dif-
ference in the degree of variation exhibited by bold and
shy individuals.
Methods
This study was conducted on a population of gray mouse
lemurs housed at the Duke Lemur Center (DLC) in
Durham, NC. A total of 14 individuals were sampled,
ranging in age from 1 to 8 years old. Age classes were
assigned as follows: Class 1 ([1 and \3 years;
n
males = 4, nfemales = 4), Class 2 ([3 and \7 years;
n n
males = 1, females = 1), Class 3 ([7 years;
n n
males = 2, females = 2). All object and food trials
were recorded using a Digital8 video camera (Sony DCR-
TRV280) and lemur behavioral responses to trials were
quantified from the tapes by JLV. A data table containing
information on individuals and responses is available in
Dryad, http://dx.doi.org/10.5061/dryad.5m083.
123
Primates
Behavioral experiments
Novel object assessment
Individual were tested for their willingness to explore a novel
object. Each individual was tested alone in his or her own
enclosure. Individuals were tested for their responses to both
control and novel objects. Control objects were chosen from
items the DLC staff uses for enrichment, thus representing
objects individuals were familiar with (e.g. tissue box, yogurt
container, toilet paper roll). Novel objects were chosen that
contained a shape, color, or texture feature the animals were
unfamiliar with. Items used included a miniature wooden
chair, a wooden heart, a wooden ladybug, plastic keys, an egg
carton, an orange ball, or a stuffed toy frog. All novel objects
were confirmed as novel by DLC staff (Dr. Sarah Zehr,
personal communication) and approved for use by a DLC
veterinarian to ensure animal safety. Each trial consisted of
four parts as follows: (1) a 5-min observation period with the
observer in the room, (2) presentation of either a control
object or novel object in the center of the enclosure, (3) a
5-min observation period, and (4) the presentation of either a
control or novel object in the center of the enclosure. Control
and novel objects were selected prior to the start of each trial
using a random number generator and were presented in
random order by flipping a coin. Each mouse lemur was
given 5 min to explore the object. The individual measures
of boldness we quantified were (a) latency to approach the
object, (b) how many times the object was approached,
(c) how many times it was touched, (d) how many times the
object was handled, and (e) the total time spent near or
handling the object. ‘Approached’ was defined as being close
enough to the object to touch it. After 5 min, the object was
removed and following a 5-min observation period, the
second object was placed in the enclosure for 5 min. Mouse
lemurs that did not approach an object, whether control or
novel, during the 5-min experimental period were assigned a
score of 5 min. A second trial was repeated 1 week later.
Novel food assessment
At least 1 week after both novel item trials were com-
pleted, the mouse lemurs were tested in their enclosures for
their inclination to consume a control and novel food item.
Following the same procedure detailed above, individuals
were presented with a control or novel food item for 5 min.
Food was presented in a stainless steel food dish prior to
the mouse lemurs receiving their afternoon meal. Control
food items were apples or bananas and the novel foods,
after being deemed safe by the DLC veterinarian, were
mango and papaya. The novel fruits were chosen based on
recommendations from DLC staff as having a high prob-
ability of being perceived as novel (Dr. Sarah Zehr,
Primates

personal communication). The individual measure of Table 1 Correlation matrix of novel object variables measured
boldness we quantified was the latency to grab and con- where V1 latency to approach the object, V2 number of times the
object was approached, V3 number of times the object was touched,
sume the food item. All individuals that grabbed the food
V4 number of times the object was handled, and V5 total time spent
consumed it. If the food was not touched during the near or handling the object
experiment, individuals were given scores of 5 min. A
Novel object V1 V2 V3 V4 V5 PC1 Eigen
second trial was repeated 1 week later. variables value

V1 1.0 – – – 0.63 3.08

Response to handling
DLC staff regularly caught and handled most individual
mouse lemurs in this study on a weekly basis for the pur-
poses of cleaning enclosures, routine weighing, or taking
breeding season measurements. An agitation score was
given to each mouse lemur based on behavior during
handling. DLC staff recorded the behaviors of individuals
on a check sheet with defined categories, provided by JLV.
We attributed 0 points if the mouse lemur was willingly
handled, 1 point if he/she vocalized, 1 point for urination, 1
point for defecation, 2 points for struggling, and 3 points
for biting. Therefore, we used an agitation index where an
individual could score a minimum of 0 (no agitation) and a
maximum of 8 (high agitation). Between 1 and 18 inde-
pendently collected scores were obtained for each indi-
vidual (X ± SE = 9.43 ± 1.69; N = 14). Because some
individuals were handled more frequently and males may
have been handled more than females for the collection of
breeding season measurements, we tested for any bias due
to sex, age-class, and the number of times that an indi-
vidual was handled. There was no relationship an indi-
vidual’s agitation index and sampling effort, sex, age class,
or sex-age class interaction (F4,9 = 1.21, P = 0.37).
Statistical analyses
We performed principal component analysis (PCA) on the
novel object behavioral measures to see if we could reduce
these variables and we also determined whether there were
any correlations among the variables (Table 1). Next, we
compared individual behavioral responses across experi-
ments and agitation indices to assess individual consistency
across the different measures. Across-context correlations
were calculated using the nonparametric Spearman’s rank
correlation test. Only individuals for whom data were
available were used in each comparison. One individual died
and one individual could not be tested due to logistical issues
prior to completion of the object and food trials, and one
individual was only handled once and was excluded from
tests involving handling (see Dryad, http://dx.doi.org/10.50
61/dryad.5m083).
We were also interested in quantifying behavioral
repeatability (r) in the mouse lemurs. To do so, we cal-
culated repeatability scores to determine how consistent
V2 -0.79 1.0 – –
V3 -0.49 0.69 1.0 –
V4 -0.36 0.17 -0.19 1.0 –
V5 -0.54 0.90 0.70 -0.21 1.0
PC1 reflects the only PCA loading for novel objects that had an
eigenvalue greater than 1
individuals were within each behavioral measure by fol-
lowing the protocol described in Lessells and Boag (1987),
where (r) represents the within-class correlation coeffi-
cient. Where necessary, data were normalized by log
transformation. We corrected for unequal sample sizes
when calculating the repeatability of handling by using the
weighted average number of observations (Gelman and
Hill 2007). We determined significance of repeatability by
calculating the 95 % confidence intervals (CIs) for each
measure as outlined in Nakagawa and Schielzeth (2010),
where significance was based on whether the 95 % CIs
overlapped 0. Lastly, because individuals were handled
multiple times and agitation indices were normally dis-
tributed, we used generalized linear models (GLM) to
determine whether individuals with higher agitation scores
showed less variation in their behavioral response using the
coefficient of variation (CV) as the dependent variable.
Sample sizes were relatively small and although multiple
tests were performed, we did not make Bonferroni cor-
rections due to the inherent low statistical power (see
Nakagawa 2004).
Results
Following PCA, in the novel object test only the first
component had an eigenvalue greater than 1, which
explained 63 % of the total variance (Table 1). Given the
strong correlation among the measures and the similar
results whether we used only the latency to approach or
PC1, we decided to use only the latency to approach in
further analyses. As expected, the small population of
captive mouse lemurs showed variation in mean (±SE)
agitation index per individual, suggesting behavioral
response differences among individuals (see Dryad,
http://dx.doi.org/10.5061/dryad.5m083). The Spearman’s
rank correlation between agitation index and the latency to

123

Mean latency to approach (sec)
300
225
150
75
0
0 1 2 3 4 5 6 7
Mean agitation index
Fig. 1 Relationship between response to handling and the latency to
approach a novel object for mouse lemurs. The relationship was tested
using the nonparametric Spearman’s correlation: rs = 0.63,
P = 0.03, N = 12
2
1.5
CV
1
0.5
0
0 2 4 6
Agitation index
Fig. 2 Relationship between response to handling (agitation index)
and the log coefficient of variation of the agitation index (GLM:
r2 = 0.77, F1,11 = 37.85, P = 0.001)
approach a novel object was positively correlated
(rs = 0.63, P = 0.03; Fig. 1). A similar result was found
when examining the relationship between agitation index
and the latency to approach a control object (rs = 0.74,
P = 0.0006), suggesting that individuals showing greater
agitation during handling took longer to explore any object
placed in a novel location in their enclosure. However,
neither agitation index nor latency to approach a novel
object was significantly correlated with the latency to
consume a novel food (handling score: rs = 0.13,
P = 0.71; object: rs = 0.36, P \ 0.29). Agitation index
was also not significantly correlated with the latency to
consume a control food (rs = 0.06, P = 0.85). In addition,
the latency to approach a novel object was not significantly
correlated with the latency to consume a control food
(rs = 0.27, P = 0.43), nor was the latency to approach a
control object and consume a novel food (rs = 0.47,
P = 0.15).
We measured repeatability for agitation index, latency
to approach the novel object, and latency to consume the
123
Primates
novel food. Of these three behavioral traits, both the agi-
tation index (r = 0.73, 95 % CI 0.57–0.89) and latency to
approach the novel object (r = 0.36, 95 % CI 0.20–0.52)
were repeatable, but the latency to consume the novel food
was not significantly repeatable (r = 0.22, 95 % CI -0.10
to 0.54). Lastly, higher agitation index was negatively
correlated with the coefficient of variation in agitation
index (r2 = 0.77, F1,11 = 37.85, P = 0.001), suggesting
these individuals showed less variation in their response
(Fig. 2).
8
Discussion
We tested the hypotheses that individuals varied in their
responses to behavioral tests, that individuals were con-
sistent in their responses, and that there were correlations
among responses to the different tests. Results reveal
considerable and repeatable between-individual variation
in two of the three behavioral tests we administered.
Individuals showed consistent differences in their response
to handling and exploratory behavior (control and novel
object). Although individuals with higher agitation scores
also were less exploratory, these patterns were not con-
sistent with the novel food item test. The relationship
between handling and investigation of objects was signif-
8 icant, regardless of whether the object was novel or
familiar. This suggests that the novelty of placement in the
enclosure may be the stimulus individuals were responding
to. Regardless of whether a control object is used or not,
one cannot decouple the object from novel placement.
Therefore, we suggest that individuals varied in their
willingness to explore objects placed in novel locations in
their enclosures. This variation was correlated with han-
dling such that individuals with a decreased latency to
explore objects were also less agitated when being handled.
These findings are consistent with what has been charac-
terized as boldness in other species (Wilson and Godin
2009). Although a general relationship has been reported
between conspecific aggression and boldness, our results
suggest that agitation during human handling may reflect
shyness, or anxiety as suggested by Carter et al. (2012).
This is in contrast to results reported by Réale et al. (2000),
where aggression in response to human handling was
correlated with measures of boldness in bighorn ewes, Ovis
Canadensis. This difference may reveal important species
variability in behavioral responses that should be
considered.
Despite the lack of correlation between the novel food
test and the other measures, we do not characterize this as a
lack of behavioral syndromes per se in mouse lemurs.
Rather, there are two possibilities. First, perhaps the food
items chosen as novel may not necessarily have been
Primates
perceived as such by individuals. While individuals were
not fed those particular fruits prior to these experiments, it
is possible that there was some feature familiar to the
mouse lemurs (e.g., smell, consistency, presentation in
familiar food dish). Second, recent studies of wild baboons,
Papio ursinus, revealed a lack of correlation among novel
food and an individual’s threat response (Carter et al.
2012). Such findings have also been reported for other
species (e.g., pumpkinseed sunfish, Lipomis gibbosus;
Coleman and Wilson 1998).
This study also addresses two key areas of current ani-
mal personality research: repeatability and flexibility. In
addition to the correlation between handling and novel
object, we found significant repeatability within individuals
across those two contexts. This strengthens the evidence
for behavioral correlations in mouse lemurs reported
for wild populations (Dammhahn 2012; Dammhahn and
Almeling 2012). The lack of repeatability in the novel food
test is also consistent with its lack of relationship with the
other two experiments and supports our suspicion that we
were unable to account for how the animals may have
actually perceived the food items or that novel foods
generally do not provide a good assay for personality tests
(see Carter et al. 2012).
In addition, the finding that individuals who were more
agitated in response to handling exhibited less variation in
that response may have implications for how flexible dif-
ferent personality types are in their behavioral responses. If
we assume that greater agitation and a lower willingness to
explore a novel object reflects a shy personality, then it is
possible that, in mouse lemurs, shyer individuals are also
less variable in their behavioral responses. This can have
important ecological and evolutionary consequences. For
example, shyer individuals may be less able to decouple
behavioral traits across contexts, constraining their ability
to adapt to uncertain or changing environmental conditions
(Dall et al. 2004; Sih et al. 2004a; Wilson and Godin 2009).
Alternatively, it has been suggested that bold individuals
exhibit less variation in behavioral traits across a bold-shy
axis (decreased behavioral plasticity) in comparison to
shyer individuals (Dochtermann and Jenkins 2007). Such
decreased variation has been observed in behaviors such as
neophobia and the likelihood of forming routines (Sih et al.
2004a; Dochtermann and Jenkins 2007). However, consis-
tent with other species, here the opposite pattern was
observed, with bolder individuals showing greater variation
or flexibility in behavioral responses than shyer individuals
(Rodriguez-Prieto et al. 2011; Dingemanse et al. 2012). This
suggests important species-specific differences that warrant
further study. As behavioral syndrome studies are emerging
as a vital part of behavioral ecology, results of such studies
have important implications for animal welfare and conser-
vation practices. A better understanding of individual
variation in behavior can be used to improve conditions for
captive animals and potentially improve the success of ani-
mal reintroduction back into the wild (Bremner-Harrison
et al. 2004; Powell and Svoke 2008; Powell 2010; Powell and
Gartner 2011). However, the effect of behavioral syndromes
and plasticity for different behavioral types, which have
implications for how well individuals cope with changing
conditions, remain unclear.
Acknowledgments Our study was approved by the institutional
Animal Care Committee at Duke University (A-227-09-08) and
adheres to the guidelines of the American Society of Primatologists’
Principles for the Ethical Treatment of Non-Human Primates. We
thank the DLC staff for their assistance with handling data collection
and other logistical support. We thank Dr. Sarah Zehr and two
anonymous reviewers for comments that greatly improved this
manuscript.
References
Bell AM (2005) Differences between individuals and populations of
three-spined stickleback. J Evol Biol 18:464–473
Bell AM, Stamps JA (2004) Development of behavioural differences
between individuals and populations of sticklebacks, gasteros-
teus aculeatus. Anim Behav 68:1339–1348
Biro PA, Stamps JA (2008) Are animal personality traits linked to
life-history productivity? TREE 23:361–368
Bremner-Harrison S, Prodohl PA, Elwood RW (2004) Behavioural
trait assessment as a release criterion: boldness predicts early
death in a reintroduction programme of captive-bred swift fox
(Vulpes velox). Anim Conserv 7:313–320
Carter AJ, Marshall HH, Heinsohn R, Colinshaw G (2012) How not to
measure boldness: novel object and antipredator responses are
not the same in wild baboons. Anim Behav 84:603–609
Coleman K, Wilson DS (1998) Shyness and boldness in pumpkinseed
fish: individual differences are context specific. Anim Behav
56:927–936
Dall SRX, Houston AI, Mcnamara JM (2004) The behavioral ecology
of personality: consistent individual differences from an adaptive
perspective. Ecol Lett 7:734–739
Dammhahn M (2012) Are personality differences in a small
iteroparous mammal maintained by a life-history trade-off? Proc
R Soc B 279:2645–2651. doi:10.1098/rspb.2012.0212
Dammhahn M, Almeling L (2012) Is risk taking during foraging a
personality trait? A field test for cross-context consistency in
boldness. Anim Behav 84:1131–1139
Dingemanse NJ, Réale D (2005) Natural selection and animal
personality. Behav 142:1165–1190
Dingemanse NJ, Bouwman KM, van de Pol M, van Overveld T,
Patrick SC, Matthysen E, Quinn JL (2012) Variation in
personality and behavioural plasticity across four populations
of the great tit parus major. J Anim Ecol 81:116–126
Dochtermann NA, Jenkins SH (2007) Behavioural syndromes in
Merriam’s kangaroo rats (dipodomys merriami): a test of
competing hypotheses. Proc R Soc B 274:2343–2349
Drea CM, Wallen K (1999) Low-status monkeys ‘‘Play dumb’’ when
learning in mixed social groups. Proc Natl Acad Sci USA
96:12965–12969
Gelman A, Hill J (2007) Data analysis using regression and
multilevel/hierarchical models. Cambridge University Press,
Cambridge
Gosling SD (2001) From mice to men: what can we learn about
personality from animal research? Psych Bull 127:45–86
123

Gosling SD, John OP (1999) Personality dimensions in nonhuman
animals: a cross-species review. Curr Dir Psych Res 8:69–75
Hamede RK et al (2009) Contact networks in a wild Tasmanian devil
(Sarcophilus harrisii) population: using social network analysis
to reveal seasonal variability in social behaviour and its
implications for transmission of devil facial tumour disease.
Ecol Lett 12:1147–1157
Lessells CM, Boag PT (1987) Unrepeatable repeatabilities: a common
mistake. Auk 104:116–121
Martin RD (1972) Adaptive radiation and behaviour of the Malagasy
lemurs. Phil Trans R Soc 264:295–352
Mittermeier RA, Tattersall I, Konstant WR, Meyers DM, Mast RB
(1994) Lemurs of Madagascar. Conservation International,
Washington, D.C
Möeller AP, Garamszegi LZ (2012) Between individual variation in
risk-taking behavior and its life history consequences. Behav
Ecol 23:843–853
Möeller AP, Nielson JT (2010) Fear screams and adaptations to avoid
imminent death: effects of genetic variation and predation. Ethol
Ecol Evol 22:1–20
Moretz JA, Martins ELP, Robison BD (2007) Behavioral syndromes
and the evolution of correlated behavior in zebrafish. Behav Ecol
18:556–562
Nakagawa S (2004) A farewell to Bonferroni: the problems of low
statistical power and publication bias. Behav Ecol 15:1044–1045
Nakagawa S, Schielzeth H (2010) Repeatability for Gaussian and
non-Gaussian data: a practical guide for biologists. Biol Rev
85:935–956
Pinter-Wollman N (2012) Personality in social insects: how does
worker personality determine colony personality? Curr Zool
58:579–587
Pintor LM, Sih A, Bauer ML (2008) Differences in aggression,
activity and boldness between native and introduced populations
of an invasive crayfish. Oikos 117:1629–1636
Powell DM (2010) A framework for introduction and socialization
processes in mammals. In: Kleiman DG, Thompson KV, Baer
CK (eds) Wild mammals in captivity: principles and techniques.
University of Chicago Press, Chicago
123
Primates
Powell DM, Gartner MC (2011) Applications of personality to the
management and conservation of nonhuman animals. In: Inoue-
Murayama M, Kawamura S, Weiss A (eds) From genes to
behavior: social structure, personalities, communication.
Springer, Berlin, pp 185–199
Powell DM, Svoke JT (2008) Novel environmental enrichment may
provide a tool for rapid assessment of animal personality: a case
study with giant pandas (Ailuropoda melanoleuca). J Appl Anim
Welf Sci 11:301–318
Quinn JL, Cresswell W (2005) Personality, anti-predation behaviour
and behavioural plasticity in the chaffinch Fringilla coelebs.
Behaviour 142:1377–1402
Radespiel U (2000) Sociality in the gray mouse lemur (Microcebus
murinus) in northwestern Madagascar. Am J Primatol 51:21–40
Réale D, Gallant BY, Leblanc M, Festa-Bianchet M (2000) Consis-
tency of temperament in bighorn ewes and correlates with
behaviour and life history. Anim Behav 60:589–597
Rodriguez-Prieto I, Martı́n J, Fernández-Juricic E (2011) Individual
variation in behavioral plasticity: direct and indirect effects of
boldness, exploration and sociability on habituation to predators
in lizards. Proc R Soc B 278:266–273
Sih A, Bell A, Johnson JC (2004a) Behavioral syndromes: an
ecological and evolutionary overview. TREE 19:372–378
Sih A, Bell A, Johnson JC (2004b) Behavioral syndromes: an
integrated overview. Quart Rev Biol 79:241–277
Wilson DS (1998) Adaptive individual differences within single
populations. Phil Trans Roy Soc B 353:199–205
Wilson DS (2007) Evolution for everyone: how Darwin’s theory can
change the way we think about our lives. Delacorte Press, New
York
Wilson ADM, Godin JJ (2009) Boldness and behavioral syndromes
in the bluegill sunfish, Lepomis macrochirus. Behav Ecol 20:
231–237
Wilson DS, Clark AB, Coleman K, Dearstyne T (1994) Shyness and
boldness in humans and other animals. TREE 9:442–446
Wolf M, Van Doorn GS, Leimar O, Weissing FJ (2007) Life-history
trade-offs favour the evolution of animal personalities. Nature
447:581–584