The unsolved challenge to phylogenetic correlation tests for categorical characters
Abstract
When comparative biologists observe that animal species living in caves also tend to have reduced eyes, they may see such correlation as evidence that the traits are adaptively or functionally linked: for instance, selection to maintain eye function is relaxed when light is unavailable. Such cross-species correlations cannot give definitive tests of evolutionary mechanisms, but nonetheless offer important insights into biological relationships among traits in realms as diverse as ecology (e.g., Paradis et al. 1998; Purvis et al. 2000) and genomics (e.g., von Mering et al. 2002; Barker and Pagel 2005). However, the last few decades have taught us that among-species correlative tests should take into account evolutionary relationships (Felsenstein 1985; Ridley 1989; Harvey and Pagel 1991). If phylogeny is not taken into account, an interpreted correlation may have a trivial explanation different from the biological relationship we claim. There is a correlation among species in the distribution of fur and bones in the middle ear—species with fur also have three bones in the middle ear, and vice versa. These two traits are characteristics of mammals, and absent outside the mammals. Using their shared distribution as evidence of an interesting biological relationship between fur and middle ear bones would be considered a mistake, however, for reasons understood long ago by Darwin (1872): We may often falsely attribute to correlated variation structures which are common to whole gr…
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Maddison and FitzJohn (2015) simulated comparative data and reported a preponderance of significant results using Pagel's correlation test (1994) and Maddison's (1990) concentrated changes test.
“In their article, Maddison and FitzJohn (2015) simulated comparative data and reported a preponderance of significant results using Pagel's correlation test (1994) and Maddison's (1990) concentrated changes test.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
Common statistical tests for the evolutionary correlation of discrete characters are prone to reporting a significant association even when the pattern is driven by a single or very few independent transitions.
“Maddison and FitzJohn (2015) recently demonstrated that common statistical tests (e.g., Maddison 1990;Pagel 1994) for the evolutionary correlation of discrete characters are prone to reporting a significant association even when the pattern is driven by a single (or, very few) independent transition(s) from one character state to another.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
These scenarios ultimately reduce to whether or not coincident unreplicated events are evidence of a causal link between traits.
“As with Case Study I, these scenarios ultimately reduce to whether or not coincident unreplicated events are evidence of a causal link between traits (Maddison and FitzJohn 2015), a problem we will address in Case Study III.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn 2015
Maddison and FitzJohn (2015) demonstrated that commonly used methods return significant correlations all the time, in scenarios that defy statistical intuition.
“As we mention above, Maddison and FitzJohn (2015) recently demonstrated that commonly used methods return significant correlations all the time—and in scenarios that seem to defy our statistical intuition.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
This is a clear case of phylogenetic pseudoreplication.
“This seems to be a clear case of phylogenetic pseudoreplication (Read and Nee 1995;Maddison and FitzJohn 2015).”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn 2015
The authors ultimately disagree with Maddison and FitzJohn (2015) that finding whether two variables are part of the same adaptive/functional network constitutes a weak conclusion; the challenges of making these inferences from any comparative data set are significant.
“We ultimately disagree with them that this constitutes a weak conclusion; the challenges of making these inferences from any comparative data set are significant.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
Maddison and FitzJohn (2015) hinted that the coincident occurrence of single events could be a way of measuring the evidence for a correlation, but did not work out the details as the authors have done here.
“Maddison and FitzJohn (2015) hinted that the coincident occurrence of single events could be a way of measuring the evidence for a correlation, but did not work out the details as we have done here.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
The traits used in comparative analyses are not chosen independently with respect to their phylogenetic distribution.
“As Maddison and FitzJohn (2015) point out, the traits we use in comparative analyses are not chosen independently with respect to their phylogenetic distribution (as we assumed in our analysis).”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
The authors agree with Maddison and FitzJohn (2015) that ascertainment bias is likely prevalent in empirical studies.
“We agree with Maddison and FitzJohn (2015) that this type of ascertainment bias is likely prevalent in empirical studies, even if it is usually more subtle than testing for a correlation between milk and middle ear bones.”
From Rethinking phylogenetic comparative methods (2018) · cites this paper as Maddison and FitzJohn (2015)
The level of significance (the size of a P-value) will depend on the size of the effect and the number of data points, tested against a null hypothesis of exactly zero difference, or exactly zero correlation.
“Usually, the level of significance (the size of a P-value) will depend on the size of the effect and the number of data points, tested against a null hypothesis of exactly zero difference, or exactly zero correlation119–121.”
From A phylogenetic approach to comparative genomics (2025) · cites this paper as [119]
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Rethinking phylogenetic comparative methods
Uyeda, J. C., Zenil-Ferguson, R., & Pennell, M. W. (2018) · Systematic Biology · doi:10.1093/sysbio/syy031 · Cited by 313 · see citing works
Ritual human sacrifice promoted and sustained the evolution of stratified societies
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A phylogenetic approach to comparative genomics
Dewar, A. E., Belcher, L. J., & West, S. A. (2025) · Nature Reviews Genetics · doi:10.1038/s41576-024-00803-0 · Cited by 41 · see citing works
A single loss of photosynthesis in the diatom order Bacillariales (Bacillariophyta)
Onyshchenko, A., Ruck, E. C., Nakov, T., & Alverson, A. J. (2019) · American Journal of Botany · doi:10.1002/ajb2.1267 · Cited by 18 · see citing works
- Authors
- Maddison, W. P., & FitzJohn, R. G.
- Affiliations
- University of British Columbia (CA)
- Corresponding author
- Maddison, Wayne P.
- Venue
- Systematic Biology
- Publisher
- Oxford University Press (OUP)
- Published
- 2014-09-09
- Volume
- 64
- Issue
- 1
- Pages
- 127-136
- ISSN
- 1063-5157 (ISSN-L)
- 1076-836X
- OpenAlex
- W2086293112
- Cited by
- 468
- References
- 2
- Citations by year
- 2026: 33 · 2025: 32 · 2024: 31 · 2023: 35 · 2022: 34 · 2021: 32 · 2020: 31 · 2019: 45 · 2018: 54 · 2017: 47 · 2016: 55 · 2015: 34 · 2014: 4
- FWCI
- 31.27
- Type
- journal article
- Language
- en
- Open access
- closed
- Locations
- Systematic Biology (publishedVersion, public-domain)
- PubMed (publishedVersion)
- Topics
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
- Physical Sciences › Mathematics › Geometry and Topology › Morphological variations and asymmetry
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Subterranean biodiversity and taxonomy
- Keywords
- pseudoreplication, phylogenetic signal, trait correlation, comparative analysis, phylogenetic signal strength
- MeSH terms
- Animals, Classification, Models, Biological, Phylogeny, Biodiversity, Genetic Speciation
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