Oops. Sorry the citation is Schluter, Price, Mooers, Ludwig 1997. Likelihood of 
ancestor states in adaptive radiation. Evolution 51:1699-1711.  This issue has 
been known for a long time. 


On Jul 30, 2013, at 6:51 AM, Marguerite Butler <mbutler...@gmail.com> wrote:

> Hi Tom,
> 
> One thing to keep in mind is the information content of the data relative to 
> what you are trying to infer. Basically, you have data only at the tips, but 
> are trying to infer the state of the root deep in the tree. So therefore 
> there is actually very little information being brought to bear on this 
> problem. In this case, whatever answer you get will very strongly reflect the 
> assumptions of the model that you apply and the structure of the tree. Put 
> another way, if you were to construct error bars around this character state 
> estimate, you would see that they are huge (See Moers et al. 1997 in 
> Evolution).   
> 
> It sounds like you are expecting a linear parsimony reconstruction. Why not 
> just use that? Your character does not change very much on the tree. This is 
> basically what your ML answer is telling you anyway, more than 50% chance of 
> red at the base.  
> 
> Marguerite
> 
> On Jul 30, 2013, at 4:38 AM, Liam J. Revell <liam.rev...@umb.edu> wrote:
> 
>> Hi Tom.
>> 
>> There is no reason to expect that the marginal ancestral state 
>> reconstructions at the root node (empirical Bayesian posterior 
>> probabilities) should match your tip frequencies or prior probabilities. 
>> Imagine the following scenario: you have one diverse clade comprising 50% of 
>> extant taxa that all diverged recently from a common ancestor share state 
>> "B"; whereas state "A" is found in all the other tips of the tree, some of 
>> which are in clades originating near the root. We would not expect posterior 
>> probabilities at the root node to mach the empirical frequencies of our 
>> state at the tips (50:50). In fact, we might expect that our reconstructed 
>> state at the root of the tree would be strongly "A".
>> 
>> In your specific case, state "0" is found in three clades that originate 
>> nearer to the root, whereas more nested clades are exclusively in state "1". 
>> This is why - in spite of its relative rarity across the tips of the tree - 
>> there is still some reasonable (PP~0.3) posterior probability under the 
>> model that the root is in state "0". This is not an error that needs to be 
>> corrected - it is just what your data, model, and tree tell us about the 
>> ancestral node of the phylogeny.
>> 
>> All the best, Liam
>> 
>> Liam J. Revell, Assistant Professor of Biology
>> University of Massachusetts Boston
>> web: http://faculty.umb.edu/liam.revell/
>> email: liam.rev...@umb.edu
>> blog: http://blog.phytools.org
>> 
>> On 7/30/2013 6:10 AM, Tom Wenseleers wrote:
>>> Dear all,
>>> Many thanks for all your advice so far. I have now moved to using rayDISC 
>>> in package corHMM to reconstruct marginal maximum likelihood ancestral 
>>> state reconstructions, using the method of Maddison et al (2007) and 
>>> FitzJohn et al (2009) to fix the prior probabilities at the root (setting 
>>> it to the observed frequency at the tips doesn't change much).
>>> The code I have is
>>> 
>>> library(ape)
>>> library(corHMM)
>>> tree=read.tree("http://www.kuleuven.be/bio/ento/temp/tree.tre";)
>>> data=read.csv(file="http://www.kuleuven.be/bio/ento/temp/data.csv";)
>>> rownames(data)=data[,1]
>>> ASR=rayDISC(tree,data,ntraits=1,charnum=1,model="ER",node.states="marginal",root.p="maddfitz")
>>> plot(tree, cex=0.6, show.tip.label=TRUE, ljoin=2,lend=2,label.offset=0.02)
>>> nodelabels(pie=ASR$states,piecol=c("white","red"), cex=0.45)
>>> tiplabels(pch = 22, bg = ifelse(data[tree$tip.label, ][,2],"red","white"), 
>>> col="black",adj = c(0.51, 0.5), cex = 0.6)
>>> 
>>> I still get unusually low marginal ML values for the trait being 1 at the 
>>> basal nodes though (ca. 0.7, which is very low considering that 89% of my 
>>> species have the trait).
>>> Would anyone be able to offer advice on why one could get the reconstructed 
>>> root ML value to be so much lower than the actual observed frequency of the 
>>> trait at the tips, and what could be a solution to obtaining a more 
>>> realistic ML reconstruction? (I also tried diversitree and phangorn, but 
>>> they all give similar results)
>>> 
>>> Cheers,
>>> Tom
>>> 
>>> 
>>> 
>>> -----Original Message-----
>>> From: Jack Viljoen [mailto:javilj...@gmail.com]
>>> Sent: 30 July 2013 10:23
>>> To: Tom Wenseleers
>>> Subject: Re: [R-sig-phylo] Question on ace ML reconstruction of discrete 
>>> binary character
>>> 
>>> Hello, Tom.
>>> 
>>> I was just wondering if the higher uncertainty at the basal nodes isn't 
>>> expected, particularly given the long branches descended from them?
>>> 
>>> Since this is an ML estimate and not a Bayesian one, surely the concept of 
>>> priors does not apply? My understanding is that ace() actually only 
>>> estimates the root node and that other methods are required to properly 
>>> estimate the states at the other nodes. I'm basing this on these posts from 
>>> Liam Revell earlier this year:
>>> http://blog.phytools.org/2013/03/conditional-scaled-likelihoods-in-ace.html
>>> http://blog.phytools.org/2013/03/a-little-more-on-ancestral-state.html
>>> 
>>> I hope those links shed some light on the matter, or that someone who knows 
>>> about this stuff has responded to you off-list as well.
>>> 
>>> Good luck,
>>> Jack
>>> 
>>> 
>>>> ----------------------------------------------------------------------
>>>> 
>>>> Message: 1
>>>> Date: Mon, 29 Jul 2013 16:00:28 +0000
>>>> From: Tom Wenseleers <tom.wensele...@bio.kuleuven.be>
>>>> To: "r-sig-phylo@r-project.org" <r-sig-phylo@r-project.org>
>>>> Subject: [R-sig-phylo] Question on ace ML reconstruction of discrete
>>>>        binary  character
>>>> Message-ID:
>>>> 
>>>> <37efc97028f3e44082acc5cbec00563011294...@icts-s-mbx7.luna.kuleuven.be
>>>>> 
>>>> 
>>>> Content-Type: text/plain; charset="us-ascii"
>>>> 
>>>> Dear all,
>>>> 
>>>> I just did some ancestral state reconstructions of binary characters 
>>>> (screenshot attached) using ace (using an equal rate discrete character 
>>>> reconstruction) . Everything seems to make sense to me, except the two 
>>>> basal nodes, where I end up with quite low likelihoods for my red 
>>>> character being 1 (cf. the pie charts), even though I get higher 
>>>> likelihoods at practically all of the more shallow nodes in the tree. Any 
>>>> ideas why one can get a result like this, and what I could potentially do 
>>>> about it, since it doesn't seem quite right to me?
>>>> 
>>>> 
>>>> 
>>>> Cheers,
>>>> 
>>>> Tom
>>>> 
>>>> ______________________________________________________________________
>>>> _________________
>>>> 
>>>> Prof. Tom Wenseleers
>>>> *      Lab. of Socioecology and Social Evolution
>>>>           Dept. of Biology
>>>>           Zoological Institute
>>>>           K.U.Leuven
>>>>           Naamsestraat 59, box 2466
>>>>           B-3000 Leuven
>>>>           Belgium
>>>> * +32 (0)16 32 39 64 / +32 (0)472 40 45 96
>>>> * tom.wensele...@bio.kuleuven.be
>>>> http://bio.kuleuven.be/ento/wenseleers/twenseleers.htm
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
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>>>> ------------------------------
>>>> 
>>>> Message: 2
>>>> Date: Mon, 29 Jul 2013 21:45:53 +0000
>>>> From: Tom Wenseleers <tom.wensele...@bio.kuleuven.be>
>>>> To: "r-sig-phylo@r-project.org" <r-sig-phylo@r-project.org>
>>>> Subject: Re: [R-sig-phylo] Question on ace ML reconstruction of
>>>>        discrete        binary character
>>>> Message-ID:
>>>> 
>>>> <37efc97028f3e44082acc5cbec00563011294...@icts-s-mbx7.luna.kuleuven.be
>>>>> 
>>>> 
>>>> Content-Type: text/plain; charset="us-ascii"
>>>> 
>>>> Dear all,
>>>> @Arne: yes I think it has to do something with the priors for the root. 
>>>> I'm not sure what prior ace uses - I think equal, which in my case would 
>>>> not be so appropriate given that nearly all species have the trait. Would 
>>>> anyone know by any chance whether in ape it is possible to have ace use a 
>>>> prior for the root which would reflect the frequency at the tips, and if 
>>>> so, how one could specify this?
>>>> 
>>>> Cheers,
>>>> Tom
>>>> 
>>>> From: Arne Mooers [mailto:amoo...@sfu.ca]
>>>> Sent: 29 July 2013 20:10
>>>> To: Tom Wenseleers
>>>> Subject: Question on ace ML reconstruction of discrete binary
>>>> character
>>>> 
>>>> Hoi Tom,
>>>> 
>>>> What is the default prior on the root in ace? Different approaches use 
>>>> different priors (=observed frequency at tips, equal, equal to tested 
>>>> ratio of q's, etc.) That has had a big affect on reconstructions I have 
>>>> done in the past.
>>>> 
>>>> Cheers,
>>>> 
>>>> Arne Mooers
>>>> 
>>>> 
>>>> Begin forwarded message:
>>>> 
>>>> From: Tom Wenseleers
>>>> <tom.wensele...@bio.kuleuven.be<mailto:tom.wensele...@bio.kuleuven.be>
>>>>> 
>>>> Date: 29 July, 2013 9:00:28 AM PDT
>>>> To: "r-sig-phylo@r-project.org<mailto:r-sig-phylo@r-project.org>"
>>>> <r-sig-phylo@r-project.org<mailto:r-sig-phylo@r-project.org>>
>>>> Subject: [R-sig-phylo] Question on ace ML reconstruction of discrete
>>>> binary character
>>>> 
>>>> Dear all,
>>>> I just did some ancestral state reconstructions of binary characters 
>>>> (screenshot attached) using ace (using an equal rate discrete character 
>>>> reconstruction) . Everything seems to make sense to me, except the two 
>>>> basal nodes, where I end up with quite low likelihoods for my red 
>>>> character being 1 (cf. the pie charts), even though I get higher 
>>>> likelihoods at practically all of the more shallow nodes in the tree. Any 
>>>> ideas why one can get a result like this, and what I could potentially do 
>>>> about it, since it doesn't seem quite right to me?
>>>> 
>>>> Cheers,
>>>> Tom
>>>> 
>>>> ______________________________________________________________________
>>>> _________________
>>>> 
>>>> Prof. Tom Wenseleers
>>>> *      Lab. of Socioecology and Social Evolution
>>>>           Dept. of Biology
>>>>           Zoological Institute
>>>>           K.U.Leuven
>>>>           Naamsestraat 59, box 2466
>>>>           B-3000 Leuven
>>>>           Belgium
>>>> * +32 (0)16 32 39 64 / +32 (0)472 40 45 96
>>>> *
>>>> tom.wensele...@bio.kuleuven.be<mailto:tom.wensele...@bio.kuleuven.be>
>>>> http://bio.kuleuven.be/ento/wenseleers/twenseleers.htm
>>>> 
>>>> 
>>>> 
>>>> [cid:image001.jpg@01CE8CB5.C244A190]__________________________________
>>>> _____________ R-sig-phylo mailing list -
>>>> R-sig-phylo@r-project.org<mailto:R-sig-phylo@r-project.org>
>>>> https://stat.ethz.ch/mailman/listinfo/r-sig-phylo
>>>> Searchable archive at
>>>> http://www.mail-archive.com/r-sig-phylo@r-project.org/
>>>> 
>>>> _____________________________
>>>> Dr. Arne Mooers
>>>> Biology, Simon Fraser University
>>>> 8888 University Drive., Burnaby BC V5A 1S6 Canada
>>>> amoo...@sfu.ca<mailto:amoo...@sfu.ca>
>>>> +1 778 782 3979
>>>> skype: arnemooers
>>>> www.sfu.ca/~amooers<http://www.sfu.ca/~amooers>
>>>> www.sfu.ca/fabstar<http://www.sfu.ca/fabstar>
>>>> www.scientists-4-species.org<http://www.scientists-4-species.org>
>>>> hesp.irmacs.sfu.ca
>>>> 7billionandyou.org
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
>>>> 
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>>>> ------------------------------
>>>> 
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>>>> End of R-sig-phylo Digest, Vol 66, Issue 22
>>>> *******************************************
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