Ann. Zool. Fennici 34: 133–137 ISSN 0003-455X
Helsinki 27 May 1997 © Finnish Zoological and Botanical Publishing Board 1997
Commentary
Why some measures of fluctuating asymmetry are so
sensitive to measurement error
Mats Björklund & Juha Merilä
Björklund, M. & Merilä, J., Department of Zoology, Uppsala University, Villavägen 9,
S-752 36 Uppsala, Sweden; Present address: Merilä, J., Laboratory of Ecology and
Animal Systematics, Department of Biology, Turku University, FIN-20014, Finland
Received 26 March 1997, accepted 1 April 1997
Introduction
Fluctuating asymmetry has recently become a sub-
ject of much theoretical and empirical interest as
well as causing considerable discussion (Houle
1997, Leamy 1997, Markow & Clarke 1997, Møll-
er & Thornhill 1997ab, Palmer & Strobeck 1997,
Pomiankowski 1997, Swaddle 1997, Whitlock &
Fowler 1997). This interest has been nurtured by
results indicating that fluctuating asymmetry is
caused by stress factors operating on the devel-
opmental system (Van Valen 1962, Palmer &
Strobeck 1986, Parsons 1990, 1992), and thus may
be a potential indicator of the amount of stress
imposed upon a given population, or conversely,
the ability of individuals to cope with stress dur-
ing their ontogeny.
As all other estimators, measurements of fluc-
tuating asymmetry are affected by measurement
error (e.g. Palmer & Strobeck 1986, Swaddle et
al. 1994, Merilä & Björklund 1995). This is par-
ticularly serious in the case of fluctuating asym-
metry since, by definition, it is expected to take
on very small values. It has previously been shown
(Swaddle et al. 1994, Merilä & Björklund 1995)
that the most commonly used measure of fluctu-
ating asymmetry, i.e. the absolute difference be-
tween the sides (|R–L|), is highly sensitive to meas-
urement error. While the fact that this measure is
prone to high measurement error has been known
for some time, the reason and its possible impli-
cations have not been examined. Furthermore, on
the basis of our results we think that measure-
ment error is a very important issue that has been
largely overlooked in the discussion.
Theory
In this note, we will give an explanation based on
basic statistical theory why both the signed and
the unsigned difference measures are so sensitive
to measurement error. By doing that, we will use
the approach taken by Whitlock (1996) with a slight,
but very important modification. Generally, fluc-
tuating asymmetry (FA) is defined as random de-
viations from perfect symmetry of bilateral traits
among a set of individuals (Palmer & Strobeck
1986). Random in this case means random in re-
lation to side, either the left or the right side being
larger, while the mean on average is zero. Thus, it
is clear that FA is a population measure of indi-
vidual asymmetry. FA can be expressed as the
signed difference of the sides (L–R) or the un-
signed (absolute) difference among the sides (| L–R|).
If we assume that the measurements of each of