Evolution and Human Behavior 28 (2007) 186 192

Facial movement varies by sex and is related to attractiveness

Edward R. Morrisona,4, Lisa Gralewskib, Neill Campbellb, Ian S. Penton-Voaka
a

Department of Experimental Psychology, University of Bristol, Bristol BS8 1TU, UK

b
Department of Computer Science, University of Bristol, Bristol BS8 1TU, UK
Initial receipt 9 October 2006; final revision received 5 January 2007

Abstract
Facial movement has received little attention in studies of human attractiveness, yet dynamic displays are an important aspect of courtship
in many species. This experiment investigated whether facial movement could be used to identify sex, and whether the ease of identification
was associated with attractiveness. We removed shape cues to sex by applying movement from individual faces to a standardised facial
model. Participants were able to distinguish between male and female animations of this model at levels above chance. Furthermore, there
was a positive association between ease of sex identification and attractiveness for female, but not male, faces. Analysis of facial movement
suggested several behaviours that are more frequent in women than men (blinking, tilting, nodding, shaking, and amount of movement).
Although some of these behaviours may be cues to sex identification, none alone was directly linked to attractiveness. Our findings suggest
that feminine motion is attractive in female faces, but sexually recognisable movement has no clear influence on male attractiveness, in
agreement with work on static faces employing composite images.
D 2007 Elsevier Inc. All rights reserved.
Keywords: Facial attractiveness; Dynamic stimuli; Sex differences; Mate choice

1. Introduction
Biological approaches to mate choice hold that cues such
as symmetry (Mller & Thornhill, 1998) and sexual
dimorphism are attractive (Andersson, 1994). In humans,
studies of facial attractiveness have argued that averageness
and symmetry may signal developmental stability and
resistance to disease (Thornhill & Mller, 1997) and
heterozygosity (Gangestand & Buss, 1993) and, hence, be
considered attractive. Sexual dimorphism is attractive in
female faces, perhaps because it signals youth and fertility,
which are valuable traits in a potential mate (Perrett et al.,
1998). These structural cues may signal useful information
to prospective mates (Fink & Penton-Voak, 2002). Sexually
dimorphic characteristics in male faces are not unequivocally attractive but can be in circumstances when putative
heritable bqualityQ is paramount in importance, such as in
short-term as opposed to long-term attractiveness (Little,
Jones, Penton-Voak, Burt, & Perrett, 2002). Two recent
reviews summarise this literature admirably (Gangestad &
Scheyd, 2005; Rhodes 2006).
4 Corresponding author. Tel.: +44 0117 954 6621.
E-mail address: ed.morrison@bris.ac.uk (E.R. Morrison).
1090-5138/$ see front matter D 2007 Elsevier Inc. All rights reserved.
doi:10.1016/j.evolhumbehav.2007.01.001

Despite the wealth of studies in the area of facial

attractiveness, the general reliance on static stimuli in the
research to date is a concern. Rubenstein (2005) reported a
low correlation between the attractiveness of individual
faces presented in static and dynamic conditions (r = .19
and r = .21, p N .18), a finding that suggests that conclusions
from experiments with static faces should be treated with
caution. A static face is, in many ways, a poor stimulus,
since faces are always dynamic in real social interactions.
Facial motion is known to convey cues about identity
(Bassili, 1978; Bruce & Valentine, 1988; Knight &
Johnston, 1997; Lander, Christie, & Bruce, 1999; Pike,
Kemp, Towell, & Phillips, 1997; Thornton & Kourtzi, 2002)
and emotional expression (Bassili, 1978, 1979). Rubenstein
(2005) showed that emotion is an important cue to
attractiveness in dynamic faces, suggesting that static
images may lack important social cues relevant to attractiveness. Indeed, Riggio, Widaman, Tucker, and Salinas
(1991) used ratings of behaviour from video sequences
to argue that dynamic information was an important
component of overall attractiveness independent of facial
(static) beauty from photographs, although the study could
not specify what aspects of movement were important in
these judgements.

E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

The role of motion in facial attractiveness deserves more

research, and a biological perspective may prove useful: the
importance of dynamic behaviours in mate choice has been
demonstrated in several species. For example, female fruit
flies choose males on the basis of their ability to bdance,Q
which signals their neuromuscular condition (MaynardSmith, 1956). Likewise, male funnel-web spiders that sway
their abdomens at high frequency are more successful at
mating (Singer et al., 2000). Video editing and computer
animation techniques have allowed experimental manipulation of such dynamic behaviours while keeping other cues
constant. For example, Rowland (1995) edited video
footage of male sticklebacks to manipulate the tempo of
their courtship displays and found that females prefer
sequences that were speeded up. Computer animation has
been used to generate artificial sticklebacks in mate
preference experiments (Kunzler & Bakker, 1998). In
humans, motion capture methods have been used to animate
a standard figure in order to study motion while controlling
for variation in static cues. In one recent study, dances by
symmetrical men were rated as most attractive, suggesting
that dynamic cues can signal underlying quality independently of static cues, which were constant across all stimuli
(Brown et al., 2005).
Specifying characteristics of movement is problematic,
but sex typicality in facial movement is a potentially
identifiable trait relevant to mate choice: sex-typical traits
are linked to mating success in many species (Andersson,
1994). Some work has investigated sex differences in
human movement. For example, a meta-analysis of studies
using point-light walkers shows that people can recognise
sex from dynamic cues, with accuracy around 70% (Pollick,
Kay, Heim, & Stringer, 2005). However, there are clear
anatomical differences between men and women that
influence gait, especially the wider female pelvis. Anatomical differences on this scale are not present in the face, but,
nonetheless, some evidence suggests that facial movement
of men and women may differ: mens movement may be
more asymmetrical (Alford & Alford, 1981; Alford, 1983),
and womens, more animated (Hall, 1985). A few studies
have tried to present dynamic faces to assess whether sex
can be identified from motion alone. Berry (1991) reported
that adults and children could identify sex from point-light
displays of dynamic faces. However, she was unable to
remove all structural cues to sex (indeed, adults could
identify sex from the static faces, although not as well as in
the dynamic condition). Hill and Johnston (2001) also
explored the role of facial motion on sex identification by
using motion-capture technology to animate an androgynous head with the movement from one of four men or four
women. People could successfully identify sex in this
experiment, and nonrigid motion (movement of the internal
features, rather than larger scale head movement) was
particularly useful for doing so. The same result was found
in similar experiments with point-light animations (Hill,
Jinno, & Johnston, 2003). In these studies, several

187

animations were generated from the same actor, meaning,

that the ratings made on them are not strictly independent
and could result in pseudoreplication if the analysis does not
take this lack of independence into account.
To our knowledge, no one has investigated whether facial
movement varies in its attractiveness independently of static
cues. In this experiment, we aimed to use a different method
to replicate the experiment of Hill and Johnston (2001) by
applying male or female movement to the same face model
and also test whether recognisability of the sex of movement
was attractive. We statistically controlled for the lack of
independence in our data using hierarchical linear modelling,
which is a form of regression that does this by explicitly
modelling the variance shared by the units of analysis. In
educational research, for example, pupils from the same
school may have exam scores that are more similar than
pupils from different schools because of the effects of the
school on performance (Bryk & Raudenbush, 1992). A twolevel hierarchical linear model nests pupils (the unit of
analysis at level 1) within schools (at level 2) and, hence,
controls for the fact that pupils exam scores are not all
independent, allowing estimation of school-level effects.
This approach also allows explanatory variables to be added
at either level (e.g., pupils socioeconomic status at level 1
and whether the school was coeducational or mixed at
level 2). In our case, we nested animations (level 1) within
the identity of the actor from which they came (at level 2).
This approach also allowed us to investigate whether any
relationship between sexually identifiable motion and attractiveness applied across different actors and across different
sequences of movement within actors. Given that dynamic
cues to attractiveness are likely to be ephemeral in nature,
investigating between- and within-target effects is important.

2. Methods
2.1. Facial animations
We obtained facial motion data from 1-min video
sequences for each of five men and five women who were
discussing the same topic (their upcoming Christmas
holidays). The volunteers were all young Caucasian
undergraduate students recruited at Stirling University.
The sequences were labelled semiautomatically with 30
landmark points per video frame (1500 frames in total;
Gralewski, Campbell, Morrison, & Penton-Voak, 2006).
The landmarks were clustered around the mouth and
eyebrows since these areas are the main components of
nonrigid facial motion (Gralewski et al., 2006) and are
important for perception of identity (Hill & Johnston, 2001)
and sex (Bruce & Valentine, 1988). These points were then
used to produce a vector, which provides a shape descriptor
of the face. The face data were scale-normalised to remove
size differences due to varying camera depth.
Since we were interested only in dynamic cues, facial
motion was extracted from the scale-normalised data for

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E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

each individual and used to animate an androgynous line

face, which was generated as the average of the 10 male and
female faces in the videos, thus removing the structural cues
to sexual dimorphism. This produced 10 1-min animations
with motion specific to each individual (see Appendix A).
Each animation was then split into six consecutive 10-s
clips, resulting in a total of 60 clips. The study was
approved by the ethics committee of the Department of
Experimental Psychology, University of Bristol.
We analysed the results using a two-level hierarchical
linear model (using HLM 6.02). This model can be
understood as linear regression looking at the effect of the
sex of motion on the rated sex of each clip, controlling for
the lack of independence between movement sequences
from the same actor. For the judgments about sex, the
outcome variable was the rated sex of each clip, which was
analysed as a binomial variable with 108 trials (i.e., number
of times each stimulus was rated male out of 108). The
level 1 predictor was sex of the movement in each clip,
which was uncentered as it was dummy-coded as 0 for
female and 1 for male. Level 2 units were the identities of
the 10 people whose movement was used, and there were no
level 2 predictors. The level 2 intercept term was treated as a
random effect (because the 10 actors represent only a
sample of all people), whereas the slope term was a fixed
effect (because male and female are the only categories into
which the predictor can fall). This type of analysis was
therefore a random coefficients model, in which the
relationship of interest lay only at level 1. The presence of
units at level 2 was merely to statistically control for the
nonindependence of stimuli derived from the same actor.
Consequently, the degrees of freedom are derived from the
number of units of analysis (60), although they are not
independent, because their shared variance has been taken
into account.
For the attractiveness ratings, the same random coefficients model was used, but now, attractiveness was the
normally distributed continuous outcome variable. The
level 1 predictor was the number of times each clip was
rated male from the first experiment, treated as a random
effect. This model allows testing of two different issues:
whether those individuals whose sex is more identifiable
seem more attractive and whether movement from a
particular individual that is sexually identifiable is also
attractive. Two models were runone for female faces and
one for male. This model can be understood as linear
regression looking at the effect of ease of sex identification
on the attractiveness of each clip, controlling for the lack
of independence.

3. Sex identification
3.1. Participants and procedures
Sixty-eight female (mean age =22.6, S.D. 6.5 years,
range = 1853) female and 40 male (mean age = 27.1,

S.D. =6.3 years, range = 1840) participants took part in

the sex identification experiment. Stimuli were presented in
random order using Eprime experimental software. Participants were instructed to judge whether each animation was
male or female after viewing it once. Participants responded
using the keyboard and did not receive feedback. Participants were not told how the stimuli had been created until
they had finished the experiment.
3.2. Results
Mean accuracy was 57% (S.D. =0.5%), with no difference between male and female raters. Sex of target,
however, did influence accuracy, which was significantly
higher for male (one-sample t test vs. 50%: mean = 61%,
SD = 0.5%, t 108 = 11.1, p b.0001) than for female (onesample t test vs. 50%: mean = 54%, SD = 0.5%, t 108 = 4.3,
p b.0001) clips (paired-samples t test for the two groups:
mean difference = 7%, t 108 =5.7, p b .0001). These effects did
not interact with participant sex (i.e., men were no better or
worse than women at identifying male or female clips).
Male and female stimuli differed in the number of times
they were categorized as male (multilevel regression,
c=0.60, t 58=2.62; p=.012). Male stimuli were 1.8 times as
likely to be judged male as female stimuli (95% CI = 1.15
2.91). The model also revealed that the variation among
level 2 units was significant [v 2(9) = 156.6, p b .0001],
meaning that clips from different actors varied in their
mean ratings. This result suggests that analyses that fail to
control for nonindependence of multiple stimuli derived
from one target are confounded.

4. Attractiveness
4.1. Participants and procedures
For this task, the same clips were blocked by sex, and
participants were informed of the sex of the faces. Consequently, a different set of 28 women (mean age= 20.1,
S.D. = 1.8 years, range = 1828) and eight men (mean
age= 20.7, S.D. = 3.0 years, range = 1827; age of rater had
no effect in any analyses) viewed the clips in random order
within block. The reason for this design, as the first
experiment showed, was that people were not very accurate
at identifying sex (see Section 3.2). This design prevents
possible misclassifications of sex influencing attractiveness
judgements. If the criteria for attractiveness differ across
male and female dynamic stimuli (as they do across male
and female static stimuli), then misclassification of sex (e.g.,
erroneously thinking a male clip were female) would lead to
potentially uninterpretable attractiveness judgements. Because participants were informed of sex, they could judge
attractiveness in the context of rating opposite sex or same
sex faces. Participants rated the faces for attractiveness on a
scale of 1 (very unattractive) to 7 (very attractive). Interrater
reliability for attractiveness ratings was high (Cronbachs
a =0.84, n = 108; for men a =0.68, n = 8; for women a = 0.86,

E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

n = 28). Most participants were undergraduate students who

received course credit for participation.

189

the scatter plot of attractiveness and rated sex typicality for

each level 2 identity of both sexes.

4.2. Results
In this analysis, we operationalized the sex typicality of a
given stimulus as the number of times that the sex of the
given clip had been correctly identified by the participants
in the first task detailed above (i.e., a clip that was correctly
identified in 100/108 trials is considered to be higher in sex
typicality than a stimulus identified correctly on 70/108
occasions). For female faces, sex typicality defined in this
way predicted attractiveness (multilevel regression, c = 1.02,
t 4 =7.08, p b.0001). There was significant variation in the
error term for the level 2 intercept [v 2(4) = 15.9, p = .004],
but not for the slope [v 2(4) =3.7, p N .500], meaning that
different actors varied in their mean attractiveness, but
correct identification of sex was associated with attractiveness in the same way across actors.
For male faces, sex typicality had no effect on
attractiveness ratings (c = 0.31, t 4 = 1.60, p =.184). Neither source of level 2 variation was significant [intercept
v 2(4) =5.9, p =.204; slope v 2(4) =6.2, p =.181]. Fig. 1 shows

Fig. 1. The relation between attractiveness and correct sex recognition of

movement for female (upper) and male (lower) faces. Data are z-standardised,
and each symbol represents one actor.

5. Movement analysis
We further investigated which specific aspects of
movement might differ between men and women and,
hence, be potential cues to sex-typical motion and/or
attractiveness. To simplify the task of defining facial
movement (which, like all biological motion, has complex
spatial and temporal properties), we concentrated on five
aspects of movement that could be objectively measured:
blinks, bshakes,Q bnods,Q btilts,Q and total movement. Blinks
were defined simply as number of blinks made during each
sequence. bShakes,Q bnods,Q and total movement were
defined objectively by measuring the movement made by
the centroid (the imaginary average vector position of all
landmark points) in each animation. Graphs of these
movements over time were smoothed (filtered) and the
number of turning points counted. A shake was defined as a
movement that exceeded a left and right threshold3.8% of
the total range of horizontal movement across all individuals. Likewise, a nod was defined as movement that
exceeded an upper and lower threshold8.0% of the total
range of vertical movement. A tilt was defined as movement
of the line between the nose and forehead landmarks that
exceeded two angle thresholds (9.7% of the total range), and
total movement was simply the distance moved by the
centroid from frame to frame. These particular thresholds
were used to strike a balance between every slight
movement becoming a nod or tilt and allowing a reasonable
and realistic number of such movements to be recorded. For
example, the rate of nodding was 3.6 times every 10 s for
women and 2.0 for men; the respective rates of tilting were
2.6 and 1.4.
Each of these behaviours was more common in
women than men when entered as a normally distributed
outcome variable into a hierarchical linear model as
used in Section 3.2. Blinks (c = 1.9, t58 = 25, p =.019),
shakes (c = 1.38, t 58 = 2.4, p = .02), nods (c = 1.7,
t 58 = 2.1, p =.039), tilts (c = 0.11, t 58 = 4.1, p =.003),
and total movement (c = 1.19, t 58 = 2.9, p =.006) were all
predicted by sex (female was dummy-coded as 0 and male
as 1, hence the negative coefficients). Variation among the
level 2 units was significant for all five variables, again
suggesting that simple linear regression would be inappropriate. We tested the possibility that these cues could be
used to identify sex by entering each variable as a predictor
of the number of times each clip was classified as male by
participants in the sex identification task. Two variables
predicted number of times rated male: blinks (c = 0.14,
t 58 = 2.4, p = .04) and total movement (c = 0.23,
t 58 = 2.5, p = .03), whereas the others did not. Variation
between level 2 units was significant for these two
variables. In other words, those clips that blinked and
moved more were more likely to be rated female. This

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E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

relationship held when sex was included in the model.

Adding the numbers of nods, shakes, and tilts resulted in a
composite variable, bfacial movements,Q which was close to
predicting number of times rated male (c = 0.04,
t 58 = 2.1, p = .06) but not attractiveness when controlling
for blinking and sex.
Odds ratios indicate that the male clips blinked 87%
(95% CI 7799%) as much as female clips and moved 79%
(6497%) as much. None of the five aspects of movement
predicted attractiveness ratings. When the same analyses
were run separately for each sex, the only significant
relationship was that total movement predicted number of
times females were rated male (c = 0.30, t 58 = 2.9,
p =.046), i.e., female clips that moved more were more
likely to be identified as female.
6. Discussion
These data suggest that men and women differ in the
way they move their faces, and that people are sensitive to
these differences. Furthermore, movement that was more
often identified as female was considered more attractive
in female faces, but no such effect was evident for male
faces. The task was by no means easy, however, as
reflected in the fairly low overall accuracy, and informally,
many participants reported finding the task difficult. Our
value of 57% is quite close to the accuracies of 68% in
Berry (1991) and about 60% in Hill and Johnston (2001),
despite differences in the stimuli and presentation details.
Male animations were more easily identified than female
animations, although accuracy for both was above chance.
Overall, there was a slight tendency for participants to rate
a face as male (53%).
Although the line-drawn stimuli here are evidently not
naturalistic, raters seem able to rate them for attractiveness
in a meaningful way. Interrater reliability in attractiveness
judgements was as high as most studies of photographic
stimuli (although see Honekopp, 2006, for a discussion of
interrater reliability in attractiveness studies). A simple
interpretation of our findings could be that since the stimuli
lacked variation in all cues except motion, attractiveness
judgments are driven by sex identification as the only source
of variation, rather than as an ecologically meaningful cue to
attractiveness. The lack of such a relationship in male faces,
however, does not support this interpretation.
Our findings are strikingly similar to those from
studies of static faces, in which femininity is attractive in
female faces (Perrett, May, & Yoshikawa, 1994; Perrett
et al., 1998), but the relationship between masculinity
and attractiveness in male faces is less simple, at least with
composite faces (Rhodes, 2006). Although Rubenstein
(2005) argues that what is attractive in static faces is not
necessarily attractive in dynamic faces, we suggest that at
least one important principle is the same: sexual dimorphism
in static faces and sex typicality in dynamic faces are
attractive (for females).

In studies with static faces, masculine faces are often

perceived as signalling negative personality traits such as
coldness and dishonesty (Penton-Voak et al., 2001). Should
these personality attributions extend to dynamic properties of
faces, it may, in part, explain why sex typicality is not
attractive in our dynamic clips. Work with composite
computer graphic images has shown that womens preferences also vary over the menstrual cycle, being higher for
masculine male faces during the fertile phase (Jones et al.,
2005; Penton-Voak et al., 1999; Penton-Voak & Perrett,
2000). Preferences also vary according to mating context,
masculine faces being more attractive as short-term or extrapair partners (Perrett et al., 1998), and according to individual
traits such as self-perceived attractiveness (Little, Burt,
Penton-Voak, & Perrett, 2001). Womens preferences may
therefore represent a strategy to trade off the benefits of good
genes signalled by a masculine face with the benefits of a
prosocial partner signalled by a less masculine face. Although
data about the role of masculine facial shapes in real faces
sometimes conflict with findings from these computer
graphic studies (Rhodes, 2006), work in other modalities
and using other techniques indicates that masculinity is
not unequivocally preferred by women in all situations
(Gangestad, Simpson, Cousins, Garver-Apgar, & Christensen,
2004; Roney, Hanson, Durante, & Maestripieri, 2006).
Dynamic facial cues may be particularly important in
investigating these tradeoffs because of the potential for
more information to be conveyed. For example, facial
motion is important in personality attribution as it improves
accuracy over static cues alone (Borkenau & Liebler, 1992;
Watson, 1989) and carries particular information about
emotion (Bente, Feist, & Elder, 1996). Dynamic information
may also signal important aspects of an individuals
underlying biology, which may be relevant to attractiveness.
Static facial masculinity is related to current levels of
circulating testosterone in men (Penton-Voak & Chen,
2004), while static facial femininity is related to current
levels of circulating oestrogen in women (Law Smith et al.,
2005). Body movement may signal symmetry and hormonal
profiles in women (Brown et al., 2005; Grammer, Filova, &
Fieder, 1997; Grammer, Fink, Mller, & Thornhill, 2003).
Because people can change the way they move their faces,
whereas static cues are invariant (at least over the short
term), more temporally precise information can be signalled.
For example, men and women might be expected to alter
their facial movement in different social situations, such as
when interacting with an attractive member of the opposite
sex compared with a rival of the same sex. Our results
demonstrate a relationship between attractiveness and sex
identification both between (some individuals move more
attractively than others across episodes) and within targets
(the attractiveness of a given individual depends on transient
dynamic cues) and support the possibility of these strategic
mate choice behaviours.
Our preliminary analysis of specific aspects of facial
motion identified several sex differences. Blinking, nodding,

E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

shaking, and tilting were all more frequent in women than in

men, and the total amount of rigid movement was higher for
female clips. Some of these cues could be used to identify
sex in the absence of structural cues, although only blinking
and total movement predicted the actual ratings made. Only
total movement was related to attractiveness in female faces,
perhaps because a more animated face signals extraversion,
which is an attractive personality characteristic (Fink, Neave,
Manning, & Grammer, 2005). These differences in facial
movement echo earlier findings in both body movement and
facial expression. Frable (1987) found that women move
their bodies more than men in expressive movement.
ODonohue and Crouch (1996), for example, also showed
that women smile and gesture more than men when speaking
to a romantic partner. None of our objectively defined
behaviours predicted attractiveness in either sex. This is
perhaps not surprising: our understanding of static face
processing advanced rapidly when feature-based approaches
to understanding facial perception gave way to configural
approaches (e.g., Tanaka & Farah, 1993). In many ways, the
technique we applied here (based on counting of discrete
behaviours) is more similar in concept to a feature-based
approach than a configural approach. More sophisticated
analysis of the spatial and temporal aspects of facial motion
may be necessary to define the properties of movement that
are associated with attractiveness ratings.
Computer animation techniques allow experimental
manipulation of the multiple signals in static and dynamic
human facial cues independently. Similar techniques have
proved fruitful in teasing apart the separate elements of
animal signals in studies of species as diverse as spiders
(Uetz & Roberts, 2002), sticklebacks (Kunzler & Bakker,
2001), and Jacky dragons (Johnstone, 2006; Ord, Peters,
Evans, & Taylor, 2002). The work reported here represents a
first step towards understanding dynamic cues to attractiveness in human faces.
Acknowledgments
The authors thank Helen Chang for collecting the raw
video footage used in this study. EM is supported by an
ESRC PhD studentship.

Appendix A. Supplementary data

Supplementary data associated with this article can
be found in the online version at www.doi:10.1016/
j.evolhumbehav.2007.01.001.
References
Alford, R. D. (1983). Sex differences in lateral facial facility:
The effects of habitual emotional concealment. Neuropsychologia, 21,
567 570.
Alford, R. D., & Alford, K. F. (1981). Sex differences in asymmetry in the
facial expression of emotion. Neuropsychologia, 19, 605 608.

191

Andersson, M. (1994). Sexual selection. Princeton, NJ7 Princeton

Univ. Press.
Bassili, J. N. (1978). Facial motion in the perception of faces and of
emotional expression. Journal of Experimental Psychology: Human
Perception and Performance, 4, 373 379.
Bassili, J. N. (1979). Emotion recognition: The role of facial movement and
the relative importance of the upper and lower areas of the face. Journal
of Personality and Social Psychology, 37, 249 258.
Bente, G., Feist, A., & Elder, S. (1996). Person perception effects of
computer-simulated male and female head movement. Journal of
Nonverbal Behavior, 20, 213 228.
Berry, D. S. (1991). Child and adult sensitivity to gender information in
patterns of facial motion. Ecological Psychology, 3, 349 366.
Borkenau, P., & Liebler, A. (1992). Trait inferences: Sources of validity at
zero acquaintance. Journal of Personality and Social Psychology, 62,
645 657.
Brown, W. M., Cronk, L., Grochow, K., Jacobson, A., Liu, K., Popovic, Z.,
& Trivers, R. (2005). Dance reveals symmetry especially in young men.
Nature, 438, 1148 1150.
Bruce, V., & Valentine, T. (1988). When a nods as good as a wink: The role
of dynamic information in facial recognition. In: M. M. Gruneberg,
Morrison P. E., &, R. N. Sykes (Eds.), Practical aspects of memory:
Current research and issues (pp. 169174). Chichester, UK7 Wiley.
Bryk, A., & Raudenbush, S. (1992). Hierarchical linear models. Newbury
Park, CA7 Sage Publications.
Fink, B., Neave, N., Manning, J.T., & Grammer, K. (2005). Facial
symmetry and the dbig fiveT personality factors. Personality and
Individual Differences, 39, 523 529.
Fink, B., & Penton-Voak, I. (2002). Evolutionary psychology of
facial attractiveness. Current Directions in Psychological Science, 11,
154 158.
Frable, D. E. S. (1987). Sex-typed execution and perception of expressive
movement. Journal of Personality and Social Psychology, 53, 391 6.
Gangestad, S. W., & Buss, D. M. (1993). Pathogen prevalence and human
mate preferences. Ethology and Sociobiology, 2, 89 96.
Gangestad, S. W., & Scheyd, G. J. (2005). The evolution of human physical
attractiveness. Annual Review of Anthropology, 34, 523 548.
Gangestad, S. W., Simpson, J. A., Cousins, A. J., Garver-Apgar, C. E., &
Christensen, P. N. (2004). Womens preferences for male behavioral
displays change across the menstrual cycle. Psychological Science, 15,
203 207.
Gralewski, L., Campbell, N., Morrison, E., & Penton-Voak, I. (2006).
Analysis of facial dynamics using a tensor framework. Journal of
Multimedia, 1, 10 21.
Grammer, K., Filova, V., & Fieder, M. (1997). The communication paradox
and possible solutions: Towards a radical empiricism. In: A. Schmitt, &
K. Schafer (Eds.), New aspects of human ethology (pp. 91120). New
York7 Plenum Press.
Grammer, K., Fink, B., Mller, A. P., & Thornhill, R. (2003). Darwinian
aesthetics: Sexual selection and the biology of beauty. Biology Review,
78, 385 407.
Hill, H., & Johnston, A. (2001). Categorizing sex and identity from the
biological motion of faces. Current Biology, 11, 880 885.
Hill, H., Jinno, Y., & Johnston, A. (2003). Comparing solid-body with
point-light animations. Perception, 32, 561 566.
Hall, J. A. (1985). Nonverbal sex differences. Baltimore7 Johns Hopkins
University Press.
Honekopp, J. (2006). Once more: Is beauty in the eye of the beholder?
Relative contributions of private and shared taste to judgments of facial
attractiveness. Journal of Experimental Psychology: Human Perception
and Performance, 32, 199 209.
Johnstone, R. A. (1996). Multiple displays in animal communication:
dBackup signalsT and dmultiple messagesT. Philosophical Transactions
of Royal Society of London: Series B, 351, 329 338.
Jones, B. C., Perrett, D. I., Little, A. C., Boothroyd, L., Cornwell, R. E.,
Feinberg, D. E., Tiddeman, B. P., Whiten, S., Pitman, R. M., Hiller, S. G.,
Burt, D. M., Stirrat, M. R., Law Smith, M. J., & Moore, F. R. (2005).

192

E.R. Morrison et al. / Evolution and Human Behavior 28 (2007) 186192

Menstrual cycle, pregnancy and oral contraceptive use alter attraction to

apparent health in faces. Proceedings of the Royal Society of London,
Series B, 272, 347 354.
Knight, B., & Johnston, A. (1997). The role of movement in face
recognition. Visual Cognition, 4, 264 273.
Kunzler, R., & Bakker, T. C. M. (1998). Computer animations as a tool in
the study of mating preferences. Behaviour, 135, 1137 1159.
Kunzler, R., & Bakker, T. C. M. (2001). Female preferences for single and
combined traits in computer animated stickleback males. Behavioral
Ecology, 12, 681 685.
Lander, K., Christie, F., & Bruce, V. (1999). The role of movement in the
recognition of famous faces. Memory Cognition, 27, 974 985.
Law Smith, M. J., Perrett, D. I., Jones, B. C., Cornwell, R. E.,
Moore, F. R., Feinberg, D. R., Boothroyd, L. G., Durrani, S. J.,
Stirrat, M. R., Whiten, S., Pitman, R. M., & Hillier, S. G. (2005).
Facial appearance is a cue to oestrogen levels in women.
Proceedings of the Royal Society of London, Series B, 273,
135 140.
Little, A. C., Burt, D. M., Penton-Voak, I., & Perrett, D. I. (2001). Self
perceived attractiveness influences human female preferences for sexual
dimorphism and symmetry in male faces. Proceedings of the Royal
Society of London, Series B, 268, 1 6.
Little, A. C., Jones, B. C., Penton-Voak, I. S., Burt, D. M., & Perrett, D. I.
(2002). Partnership status and the temporal context of relationships
influence human female preferences for sexual dimorphism in male
face shape. Proceedings of the Royal Society of London, Series B
1095 1100.
Maynard Smith, J. (1956). Fertility, mating behaviour and sexual selection
in Drosophila subobscura. Journal of Genetics, 54, 261 279.
Mller, A. P., & Thornhill, R. (1998). Bilateral symmetry and sexual
selection: A meta-analysis. American Naturalist, 151, 174 192.
ODonohue, W., & Crouch, J. L. (1996). Marital therapy and gender-linked
factors in communication. Journal of Marital and Family Therapy, 22,
87 101.
Ord, T. J., Peters, R. A., Evans, C. S., & Taylor, A. J. (2002). Digital video
playback and visual communication in lizards. Animal Behaviour, 63,
879 890.
Penton-Voak, I. S., Jones, B. C., Little, A. C., Baker, S., Tiddeman, B.,
Burt, D. M., & Perrett, D. I. (2001). Symmetry, sexual dimorphism in
facial proportions and male facial attractiveness. Proceedings of the
Royal Society of London, Series B, 268, 1617 1623.
Penton-Voak, I. S., & Chen, J. Y. (2004). High salivary testosterone is
linked to masculine male facial appearance in humans. Evolution and
Human Behavior, 25, 229 241.
Penton-Voak, I. S., & Perrett, D. I. (2000). Female preference for male faces
changes cyclically: Further evidence. Evolution and Human Behavior,
21, 39 48.

Penton-Voak, I. S., Perrett, D. I., Castles, D. L., Kobayashi, T., Burt, D. M.,
Murray, L. K., & Minamisawa, R. (1999). Menstrual cycle alters face
preference. Nature, 399, 741 742.
Perrett, D. I., Lee, K. J., Penton-Voak, I. S., Rowland, D. R., Yoshikawa, S.,
Burt, D. M., Henzi, S. P., Castles, D. L., & Akamatsu, S. (1998). Effects
of sexual dimorphism on facial attractiveness. Nature, 394, 884 887
(doi:10.1038/29772).
Perrett, D. I., May, K. A., & Yoshikawa, S. (1994). Facial shape and
judgements of female attractiveness. Nature, 368, 239 242.
Pike, G. E., Kemp, R. I., Towell, N. A., & Phillips, K. C. (1997).
Recognizing moving faces: The relative contribution of motion and
perspective view information. Visual Cognition, 4, 409 437.
Pollick, F. E., Kay, J. W., Heim, K., & Stringer, R. (2005). Gender
recognition from point-light walkers. Journal of Experimental Psychology: Human Perception and Performance, 31, 1247 1265.
Rhodes, G. (2006). The evolutionary psychology of facial beauty. Annual
Review of Psychology, 57, 199 226.
Riggio, R. E., Widaman, K. F., Tucker, J. S., & Salinas, C. (1991). Beauty
is more than skin deep: Components of attractiveness. Basic and
Applied Social Psychology, 12, 423 439.
Roney, J. R., Hanson, K. N., Durante, K. M., & Maestripieri, D. (2006).
Reading mens faces: Womens mate attractiveness judgments
track mens testosterone and interest in infants. Proceedings of
the Royal Society of London Series B, Biological sciences, 273,
2169 2175.
Rowland, W. J. (1995). Do female sticklebacks care about male courtship
vigour? Manipulation of display tempo using video playback. Behaviour, 132, 951 961.
Rubenstein, A. J. (2005). Variation in perceived attractiveness. Psychological Science, 16, 759 762.
Singer, F., Riechert, S. E., Xu, H., Morris, A. W., Becker, E., Hale, J. A., &
Noureddine, M. A. (2000). Analysis of courtship success in the funnelweb spider Agenenopsis aperta. Behaviour, 137, 93 117.
Tanaka, J. W., & Farah, M. J. (1993). Parts and wholes in face recognition.
Quarterly Journal of Experimental Psychology Human Experimental
Psychology, 46, 225 245.
Thornton, I. M., & Kourtzi, Z. (2002). A matching advantage for dynamic
faces. Perception, 31, 113 132.
Thornhill, R., & Mller, A. P. (1997). Developmental stability, disease and
medicine. Biological Reviews of the Cambridge Philosophical Society,
72, 497 548.
Uetz, G. W., & Roberts, J. A. (2002). Multisensory cues and multimodal
communication in spiders: Insights from video/audio playback studies.
Brain Behaviour and Evolution, 59, 222 230.
Watson, D. (1989). Strangers ratings of the five robust personality factors:
Evidence of a surprising convergence with self-report. Journal of
Personality and Social Psychology, 57, 120 129.