Palmer, A.R. 1994. Fluctuating asymmetry analyses: A primer. p. 335-364. In: Developmental Instability: Its Origins and Evolutionary Implications. (T. Markow, Ed.). Kluwer, Dordrecht.

(download a portable-document-file (pdf) version of the original MS with figures; size of download is 201K= 35 pages).


1.0 Fluctuating asymmetry and developmental stability: A cursory overview

2.0 Terminology

2.1 Asymmetry in an individual vs pattern of asymmetry variation in a sample

2.2 Patterns vs. processes

2.3 Glossary

3.0 Indices for describing the level of FA in a sample

3.1 FA indices

3.2 Pros & cons of different FA indices

3.3 Relationships among FA indices

3.4 General recommendations regarding FA indices

4.0 Choice of traits

4.1 Pros & cons of meristic vs metrical traits

4.2 Two idiosyncrasies of meristic traits

4.3 Single vs multiple traits

4.4 Choose traits that are developmentally independent

4.5 Choose traits that exhibit 'ideal' FA

5.0 Sample sizes

6.0 Measurement error

6.1 Why is measurement error a particular concern in studies of FA?

6.2 Error in meristic traits

6.3 Error in metrical traits

6.4 Quantizing error in image analysis systems

6.5 Recommended procedure for conducting repeated measurements

6.6 Tests for the significance of FA relative to measurement error in metrical traits

6.7 Tests for the significance of FA relative to counting error in meristic traits

7.0 Directional Asymmetry

7.1 Why test for DA in studies of FA?

7.2 Tests for DA

8.0 Departures from normality (e.g. antisymmetry and skew)

8.1 Why test for departures from normality in studies of FA?

8.2 Tests for departures from normality: general comments

8.3 Tests for departures from normality in either meristic or metrical traits

8.4 Tests for departures from normality in small samples of metrical traits

9.0 What to do when traits depart from ideal FA

10.0 Size dependence of FA

10.1 Why is size-dependence a concern in studies of FA?

10.2 Tests for size dependence of FA within samples

10.3 Tests for size dependence of FA among samples

11.0 Care when conducting multiple tests

12.0 Adequate presentation of descriptive data

13.0 Significance tests for differences in FA

13.1 Between two samples

13.2 Among three or more samples

13.3 Among samples using multiple traits simultaneously

14.0 Correlations of subtle asymmetries among traits

14.1 Between parents and offspring (heritability)

14.2 Among individuals: Are some individuals more stable developmentally than others?

14.3 Among samples

15.0 Correlations between asymmetry & fitness: Can asymmetry predict mate choice?

16.0 Checklist for studies of FA

16.0 Checklist for studies of FA

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(revised Nov. 4, 2002)