Good evening,

Apologies for the delayed response.

> Contml was originally designed for inferring phylogenies from gene frequency 
> data in closely related populations, under the assumption that the mechanism 
> of change is genetic drift.  Then it is roughly possible to assume 
> independent Brownian Motions of allele frequencies, with equal variances of 
> change.
> 
> For quantitative characters all these assumptions are highly dubious.  I try 
> to point out to the questioner that they cannot assume that the characters 
> are independent, and that they change with equal variances.  The questioners 
> rarely react with enthusiasm to this.


This appears to be the issue with all proposed methods for analyzing GM data, 
save perhaps the PC-based methods. Proposed parsimony-based methods of 
accounting for inapplicability and interdependence in discrete characters (e.g. 
Brazeau et al., 2019 
<https://academic.oup.com/sysbio/article/68/4/619/5238046>; Hopkins & St. John, 
2021 <https://academic.oup.com/sysbio/article/70/6/1163/6131693>; Goloboff et 
al., 2021 <https://onlinelibrary.wiley.com/doi/full/10.1111/cla.12456>; 
Wheeler, 2023 <https://onlinelibrary.wiley.com/doi/full/10.1111/cla.12553>; 
Goloboff & De Laet, in press 
<https://onlinelibrary.wiley.com/doi/full/10.1111/cla.12564>) aren’t easily 
applicable to continuously varying data (see review by Goloboff, 2022), and 
I’ve yet to encounter a likelihoodist alternative.

> I would encourage people to try going the PCM route: getting a tree for 
> living species and using it to infer covariances of changes in characters.  I 
> have mentioned in a few places that from that one might even use the 
> resulting inference to place fossil species on a tree (perhaps Liam Revell 
> has published that suggestion too).

I’m assuming you mean Revell et al. (2015) 
<https://academic.oup.com/evolut/article/69/4/1027/6852435>. I imagine this 
procedure is easily workable when the molecular “backbone” topology is robustly 
supported, and the developmental constraints of morphometric characters are 
well-known for both extant and extinct taxa, e.g., size and shape of mammalian 
teeth. I don’t know if it’s workable for taxa without any molecular dats, 
without close modern relatives, and with non-modern-analog morphology and 
ontogeny, e.g., graptolites or vendobionts.

> I would also note that there is a program, Threshml, distributed by my lab, 
> that uses Sewall Wright's "threshold model" to model discrete character 
> evolution, and to treat both continuous and discrete characters in the same 
> analysis.

I did not know this. Thanks!

Jacqueline S. Silviria
The Last King of the Jungle

Department of Earth & Space Science
University of Washington
Seattle, WA, USA
[email protected] <mailto:[email protected]>, [email protected] 
<mailto:[email protected]>

ResearchGate profile <https://www.researchgate.net/profile/J_Silviria>
Twitter: @JSilviria

Sent from my iPhone

> On Feb 5, 2024, at 6:33 PM, Joe Felsenstein <[email protected]> wrote:
> 
> 
> Morphmet people --
> 
> I want to concur with Jacqueline Silviria and with Chris Klingenberg about 
> the difficulties of using morphometric analyses to infer phylogenies.  Just 
> to "violently agree":  This is a more general problem with quantitative 
> characters of any kind.  I often get asked how to use the 
> continuous-character maximum likelihood program Contml in my package PHYLIP 
> to infer a phylogeny from quantitative characters such as morphological 
> measurements.  Contml was originally designed for inferring phylogenies from 
> gene frequency data in closely related populations, under the assumption that 
> the mechanism of change is genetic drift.  Then it is roughly possible to 
> assume independent Brownian Motions of allele frequencies, with equal 
> variances of change.
> 
> For quantitative characters all these assumptions are highly dubious.  I try 
> to point out to the questioner that they cannot assume that the characters 
> are independent, and that they change with equal variances.  The questioners 
> rarely react with enthusiasm to this.
> 
> Of course there is a literature (Phylogenetic Comparative Methods) where we 
> have methods (which I guess we should call "Phylogenetic Comparative Methods 
> methods") that take a tree that is presumed known, or a cloud of trees from 
> bootstrapping or a sampled Bayesian posterior, and work the other way.  
> Namely they infer the covariances between changes in a multivariate Brownian 
> Motion, given the tree.
> 
> I know that in the brilliant theory of Kendall, there are theorems showing 
> that if the individual landmark (x,y) coordinates have equal and isotropic 
> i.i.d. variances, then so will the coordinates in the shape space.  But what 
> that is doing is assuming that the differences we see are all measurement 
> error.  It does not guarantee independent i.i.d. Brownian Motion when the 
> changes along the branches of the tree are themselves covarying among 
> characters and with unequal variances.
> 
> I would encourage people to try going the PCM route: getting a tree for 
> living species and using it to infer covariances of changes in characters.  I 
> have mentioned in a few places that from that one might even use the 
> resulting inference to place fossil species on a tree (perhaps Liam Revell 
> has published that suggestion too).
> 
> I would also note that there is a program, Threshml, distributed by my lab, 
> that uses Sewall Wright's "threshold model" to model discrete character 
> evolution, and to treat both continuous and discrete characters in the same 
> analysis.
> 
> Joe
> ----
> Joe Felsenstein        [email protected] <mailto:[email protected]>,   
> [email protected] <mailto:[email protected]>
>  Department of Genome Sciences and Department of Biology,
>  University of Washington, Box 355065, Seattle, WA 98195-5065 USA
> -----
>  PS: please do not use  [email protected] 
> <mailto:[email protected]>, which is an alias 
>  that some mail systems now mistake as indicating spam.
> 
> 

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