Andrea,

It is the case that one risks getting into a bit of a circularity issue when 
using traits to estimate phylogenetic relationships and then evaluating the 
evolution of those traits on that same tree. The problem is most acute when the 
traits for both objectives are the same, are entirely correlated (or nearly 
so), and when there are no other sources of data for phylogenetic estimation 
and inference.

Generally, estimating the phylogeny for extant and fossil taxa simultaneously 
-- where molecular and morphological data are obtained from extant taxa and 
morphological only from fossils) is preferable than using a fossil-only 
backbone phylogeny, as the problem you outlined can be more pronounced with the 
latter approach.

Dean

Dr. Dean C. Adams
Distinguished Professor of Evolutionary Biology
Department of Ecology, Evolution, and Organismal Biology
Iowa State University
https://faculty.sites.iastate.edu/dcadams/
phone: 515-294-3834

From: [email protected] <[email protected]> On Behalf Of 
alcardini
Sent: Thursday, February 8, 2024 12:47 AM
To: morphmet2 <[email protected]>
Subject: [MORPHMET2] related question on comparative methods using fossils

I let the experts answering Jacqueline's questions, but have a related one. 
It's not about phylogenetic inference using morphometrics.

I've reviewed a few paleoanthropological papers where the authors used a 
phylogeny that included fossil hominins as the backbone to apply comparative 
methods to cranial shape data. At least for the fossils (with few exceptions 
like Neanderthals, where ancient DNA is available), the phylogeny was a 
morphological analysis using mostly cranial data. The phylogeny was usually 
taken from the literature and, therefore, was not using the same shape 
measurements as in the study but was still based on cranial data (meristic, 
traditional morphometric etc.). My guess is that this is a mistake because the 
data (cranial morphology, even if not exactly the same variables) are not 
independent.
This is not specific to paleoanthropological data and probably a common issue 
in the analysis of fossils with no independent (like a molecular phylogeny for 
living species) information. Am I off track?

Cheers

Andrea


---------- Forwarded message ---------
From: Jacqueline Silviria 
<[email protected]<mailto:[email protected]>>
Date: Thu, 8 Feb 2024 at 01:46
Subject: Re: [MORPHMET2] Compiling metadata on all methods of phylogenetic 
inference using GM data
To: Joe Felsenstein <[email protected]<mailto:[email protected]>>
Cc: Chris Klingenberg <[email protected]<mailto:[email protected]>>, 
Joe Felsenstein <[email protected]<mailto:[email protected]>>, andrea cardini 
<[email protected]<mailto:[email protected]>>, 
[email protected]<mailto:[email protected]> 
<[email protected]<mailto:[email protected]>>

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]<mailto:[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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