Package 'hisse'

Description

Takes a sampled set of m and k fossils from GetFossils and them as points to a plot of the phylogeny from which they were sampled.

Usage

AddFossilPoints(phy, f, ...)

Arguments

Author(s)

Jeremy M. Beaulieu

References

Vascancelos, T, B.C. O'Meara, and J.M. Beaulieu. In prep.

Description

Takes a sampled set of stratigraphic intervals from GeteStratigraphicIntervals and adds them as segments to a plot of the phylogeny from which they were sampled.

Usage

AddStratIntervals(phy, f, ...)

Arguments

Author(s)

Jeremy M. Beaulieu

References

Vascancelos, T, B.C. O'Meara, and J.M. Beaulieu. In prep.

Description

Computes the Akaike model weights for a list of HiSSE, MiSSE, and/or GeoHiSSE model fits.

Usage

GetAICWeights(hisse.results, criterion="AIC")

Arguments

Details

Function computes the model weight from their AIC values using the formula:

“delta <- mod.AIC - min( mod.AIC )”

“AICw <- exp( -0.5 * delta) / sum( exp( -0.5 * delta) )”

Function will return vector of weights

Author(s)

Daniel Caetano

Description

Sets up a set of MiSSE models (Missing State Speciation and Extinction) on a phylogeny, varying the number of parameters for turnover and extinction fraction.

Usage

generateMiSSEGreedyCombinations(max.param=52, turnover.tries=sequence(26), 
eps.tries=sequence(26), fixed.eps.tries=NA, vary.both=TRUE, shuffle.start=TRUE)

Arguments

Details

This creates the set of combinations of models to run through MiSSEGreedy as a data.frame. It has columns for the number of rates to estimate for turnover, the number of values to estimate for eps, and any fixed values for eps to apply to the whole tree. You can add your own rows or delete some to this data.frame to add more or fewer combinations. By default, this comes out ordered so that simpler models are run first in MiSSEGreedy but that is not required (but wise for most use cases), and you can reorder if you wish.

It can be worth considering how much information your tree has for diversification rates. A fully resolved tree with N taxa has 2N-2 branches and only N-1 internal node heights, which is more relevant for diversification models. So on a 50 taxon tree, there are 49 node heights – it's a bit ambitious to think you can extract, say, 5 different parameters (3 turnover rates, 1 eps value, 1 transition rate) from such a tree. People clearly try to extract more – some methods claim to use information in a tree to extract a different speciation or net diversification rate for every single tip – but that could be a tad ambitious. So setting max.param to a low value makes a lot of sense. max.param = floor(ape::Ntip(phy)/10) is ridiculously optimistic (dividing by 100 or more is probably more conservative) but if things are running well MiSSEGreedy should stop before getting to the models that are too complex.

turnover.tries and eps.tries set how many turnover and eps hidden states to try, respectively. If you wanted to try only 1, 3, and 7 hidden states for turnover you would set turnover.tries = c(1, 3, 7) for example.

Estimating extinction rates is hard. This affects all diversification models (even if all you want and look at is speciation rate, extinction rate estimates still affect what this is as they affect the likelihood). It is most noticeable in MiSSE with eps, the extinction fraction (extinction rate divided by speciation rate). One option, following Magallon & Sanderson (2001), is to set extinction fraction at set values. By default, we use theirs, 0 (meaning a Yule model - no extinction) or 0.9 (a lot of extinction, though still less than paleontoligists find). You can set your own in fixed.eps.tries. If you only want to use fixed values, and not estimate, get rid of the NA, as well. However, don't “cheat” – if you use a range of values for fixed.eps, it's basically doing a search for this, though the default AICc calculation doesn't dQuoteknow this to penalize it for another parameter.

HiSSE and thus MiSSE assume that a taxon has a particular hidden state (though they recognize that there can be uncertainty in which state it actually has). Thus, they're written to assume that we dQuotepaint these states on the tree and a given state affects both turnover and eps. So if turnover has four hidden states, eps has four hidden states. They can be constrained: the easiest way is to have, say, turnover having an independent rate for each hidden state and eps having the same rate for all the hidden states. If vary.both is set to FALSE, all models are of this sort: if turnover varies, eps is constant across all hidden states, or vice versa. Jeremy Beaulieu prefers this. If vary.both is set to TRUE, both can vary: for example, there could be five hidden states for both turnover and eps, but turnover lets each of these have a different rate, but eps only allows three values (so that eps_A and eps_D might be forced to be equal, and eps_B and eps_E might be forced to be equal). Brian O'Meara would consider allowing this, while cautioning you about the risks of too many parameters.

Value

generateMiSSEGreedyCombinations returns a data.frame to pass to MiSSEGreedy().

Author(s)

Brian C. O'Meara

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Magallon S. and Sanderson M.J. 2001. Absolute diversification rates in angiosperm clades. Evolution 55:1762-1780.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Sets up and executes a GeoHiSSE model (Hidden Geographic State Speciation and Extinction) on a very large phylogeny and character distribution.

Usage

GeoHiSSE(phy, data, f=c(1,1,1), turnover=c(1,2,3), eps=c(1,2), 
hidden.states=FALSE, trans.rate=NULL, assume.cladogenetic=TRUE, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, sann=TRUE,
sann.its=1000, bounded.search=TRUE,  max.tol=.Machine$double.eps^.50,
mag.san.start=0.5, starting.vals=NULL, turnover.upper=1000, 
eps.upper=3, trans.upper=100, restart.obj=NULL, ode.eps=0, dt.threads=1)

Arguments

Details

This function sets up and executes a more complex and faster version of the GeoHiSSE model (for the original function see GeoHisse.old). One of the main differences here is that the model allows up to 10 hidden categories, and implements a more efficient means of carrying out the branch calculation. Specifically, we break up the tree into carry out all descendent branch calculations simultaneously, combine the probabilities based on their shared ancestry, then repeat for the next set of descendent . In testing, we've found that as the number of taxa increases, the calculation becomes much more efficient. In future versions, we will likely allow for multicore processing of these calculations to further improve speed. Also, note this function has replaced the version of GeoSSE that is currently available (see GeoHisse.old).

The other main difference is that, like HiSSE, we employ a modified optimization procedure. In other words, rather than optimizing birth and death separately, GeoHisse optimizes orthogonal transformations of these variables: we let tau = birth+death define "net turnover", and we let eps = death/birth define the “extinction fraction”. This reparameterization alleviates problems associated with overfitting when birth and death are highly correlated, but both matter in explaining the diversity pattern.

To setup a model, users input vectors containing values to indicate how many free parameters are to be estimated for each of the variables in the model. This is done using the turnover and extinct.frac parameters. One needs to specify a value for each of the parameters of the model, when two parameters show the same value, then the parameters are set to be linked during the estimation of the model. For example, a GeoHiSSE model with 1 hidden area and all free parameters has turnover = 1:6. The same model with speciation rates constrained to be the same for all hidden states has turnover = c(1,2,3,1,2,3). This same format applies to extinct.frac.

Once the model is specified, the parameters can be estimated using the subplex routine (default), or use a two-step process (i.e., sann=TRUE) that first employs a stochastic simulated annealing procedure, which is later refined using the subplex routine.

The “trans.rate” input is the transition model and has an entirely different setup than speciation and extirpation rates. See TransMatMakerGeoHiSSE function for more details.

For user-specified “root.p”, you should specify the probability for each area. If you are doing a hidden model, there will be six areas: 0A, 1A, 2A, 0B, 1B, 2B. So if you wanted to say the root had to be in area 0 (widespread distribution), you would specify “root.p = c(0.5, 0, 0, 0.5, 0, 0)”. In other words, the root has a 50% chance to be in one of the areas 0A or 0B.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Value

GeoHiSSE returns an object of class geohisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Caetano, D.S., B.C. O'Meara, and J.M. Beaulieu. 2018. Hidden state models improve state-dependent diversification approaches, including biogeographic models. Evolution, 72:2308-2324.

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., W.P. Maddison, and S.P. Otto. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., P.E. Midford, and S.P. Otto. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., R.M. May, and P.H. Harvey. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Sets up and executes the original GeoHiSSE model (Hidden Geographic State Speciation and Extinction) on a phylogeny and character distribution.

Usage

GeoHiSSE.old(phy, data, f=c(1,1,1), speciation=c(1,2,3), extirpation=c(1,2), 
hidden.areas=FALSE, trans.rate=NULL, assume.cladogenetic=TRUE, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, sann=TRUE,
sann.its=1000, bounded.search=TRUE,  max.tol=.Machine$double.eps^.50,
mag.san.start=0.5, starting.vals=NULL, speciation.upper=1000, extirpation.upper=1000, 
trans.upper=100, ode.eps=0)

Arguments

Details

This function sets up and executes the original GeoHiSSE model. The model closely follows diversitree, although here we employ modified optimization procedures. As for data file format, GeoHiSSE expects a two column matrix or data frame, with the first column containing the species names and the second containing the are information. The area information need to be in the format of three numbers: 0 for area '01', 1 for area '0', and 2 for '1'. Please note that the code for the areas here differ from the make.geosse function of package diversitree. The order of the data file and the names in the “phylo” object need not be in the same order; hisse deals with this internally. Also, the character information MUST be 0, 1, or 2, otherwise, the function will return an error message.

To setup a model, users input vectors containing values to indicate how many free parameters are to be estimated for each of the variables in the model. This is done using the speciation and extirpation parameters. One needs to specify a value for each of the parameters of the model, when two parameters show the same value, then the parameters are set to be linked during the estimation of the model. For example, a GeoHiSSE model with 1 hidden area and all free parameters has speciation = 1:6. The same model with speciation rates constrained to be the same for all hidden areas has speciation = c(1,2,3,1,2,3). This same format applies to extirpation. Please note that GeoHiSSE currently works with up to 4 hidden areas. The most complex model would be speciation = 1:15 and extirpation = 1:10.

Once the model is specified, the parameters can be estimated using the subplex routine (default), or use a two-step process (i.e., sann=TRUE) that first employs a stochastic simulated annealing procedure, which is later refined using the subplex routine.

The “trans.rate” input is the transition model and has an entirely different setup than speciation and extirpation rates. See TransMatMakerGeoHiSSE.old function for more details.

For user-specified “root.p”, you should specify the probability for each area. If you are doing a hidden model, there will be six areas: 0A, 1A, 01A, 0B, 1B, 01B. So if you wanted to say the root had to be in area 0 (widespread distribution), you would specify “root.p = c(0.5, 0, 0, 0.5, 0, 0)”. In other words, the root has a 50% chance to be in one of the areas 0A or 0B.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Value

GeoHiSSE returns an object of class geohisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Caetano, D.S., B.C. O'Meara, and J.M. Beaulieu. 2018. Hidden state models improve state-dependent diversification approaches, including biogeographic models. Evolution, 72:2308-2324.

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., W.P. Maddison, and S.P. Otto. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Goldberg, E. E., L. T. Lancaster, and R. H. Ree. 2011. Phylogenetic Inference of Reciprocal Effects between Geographic Range Evolution and Diversification. Syst. Biol. 60:451-465.

Maddison W.P., P.E. Midford, and S.P. Otto. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., R.M. May, and P.H. Harvey. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Samples a simulated tree of extinct and extant lineages according to the preservation rate of the fossilized birth-death process.

Usage

GetFossils(phy, psi=0.1)

Arguments

Value

Returns a table of sampled lineages.

Description

Sets up and executes a HiSSE model (Hidden State Speciation and Extinction) on a phylogeny and character distribution.

Usage

hisse(phy, data, f=c(1,1), turnover=c(1,2), eps=c(1,2), 
hidden.states=FALSE, trans.rate=NULL, condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL, includes.fossils=FALSE, k.samples=NULL, 
strat.intervals=NULL, tip.fog=NULL, sann=TRUE, sann.its=5000, 
bounded.search=TRUE, max.tol=.Machine$double.eps^.50, starting.vals=NULL, 
fog.ip=0.01, turnover.upper=10000, eps.upper=3,trans.upper=100, 
restart.obj=NULL, ode.eps=0, dt.threads=1)

Arguments

Details

This function sets up and executes a new and faster version of the HiSSE model. Note that the four-state character-independent model can be called from this command in addition to the two-state BiSSE model and the full character-dependent HiSSE model. See vignette on how to set this up.

The “trans.rate” input is the transition model and has an entirely different setup than turnover rates and extinction fraction. See TransMatMakerHiSSE function for more details.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

For user-specified “root.p”, you should specify the probability for each state. If you are doing a hidden model, there will be four states: 0A, 1A, 0B, 1B. So if you wanted to say the root had to be state 0, you would specify “root.p = c(0.5, 0, 0.5, 0)”.

This code will completely replace the original hisse function in the next version.

Value

hisse returns an object of class hisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Examples

library(diversitree)
pars <- c(0.1, 0.2, 0.03, 0.03, 0.01, 0.01)
set.seed(4) 
phy <- tree.bisse(pars, max.t=30, x0=0)
sim.dat <- data.frame(names(phy$tip.state), phy$tip.state)

## Fit BiSSE equivalent:
trans.rates.bisse <-  TransMatMakerHiSSE(hidden.traits=0)
pp.bisse <- hisse(phy, sim.dat, hidden.states=FALSE, turnover=c(1,2), 
eps=c(1,2), trans.rate=trans.rates.bisse)

## Now fit HiSSE equivalent with a hidden state for state 1:
trans.rates.hisse <- TransMatMakerHiSSE(hidden.traits=1)
pp.hisse <- hisse(phy, sim.dat, hidden.states=TRUE, turnover=c(1,2,1,2), 
eps=c(1,2,1,2), trans.rate=trans.rates.hisse)

Description

Sets up and executes the original four state trait-independent HiSSE model (Hidden State Speciation and Extinction) on a phylogeny and character set.

Usage

hisse.null4.old(phy, data, f=c(1,1), turnover.anc=rep(c(1,2,3,4),2), 
eps.anc=rep(c(1,2,3,4),2), trans.type="equal", condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL,  output.type="turnover", sann=TRUE, 
sann.its=1000, bounded.search=TRUE, max.tol=.Machine$double.eps^.50,
starting.vals=NULL, turnover.upper=10000, eps.upper=3, trans.upper=100,
ode.eps=0)

Arguments

Details

This function sets up and executes a four-state trait independent HiSSE model. The model closely follows hisse.old. However, note that this function is no longer necessary and can be called and evaluated directly using the new hisse function.

Like hisse.old, users input vectors containing values to indicate how many free parameters are to be estimated for each of the variables in the model. However, the null-four model assumes that “turnover.anc” and “eps.anc” are linked between the two observed states. Thus, users are unlikely to alter the inputs much, aside from perhaps fixing “turnover.anc” or “eps.anc” to be equal across the four hidden states, where the “turnover.anc” input vector is set as rep(c(1,1,1,1),2). For a Yule equivalent, the input vector for “eps.anc” would be rep(c(0,0,0,0),2). For how to setup a null-two model see the example code below.

For user-specified “root.p”, you should specify the probability for each state. See help for “hisse.old” for more on other parameters for this function.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Value

hisse.null4.old returns an object of class hisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

Sets up and executes the original HiSSE model (Hidden State Speciation and Extinction) on a phylogeny and character distribution.

Usage

hisse.old(phy, data, f=c(1,1), hidden.states=TRUE, turnover.anc=c(1,1,0,0), 
eps.anc=c(1,1,0,0), trans.rate=NULL, turnover.beta=c(0,0,0,0), 
eps.beta=c(0,0,0,0), timeslice=NULL, condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL, output.type="turnover", sann=TRUE,
sann.its=1000, bounded.search=TRUE, max.tol=.Machine$double.eps^.50,
starting.vals=NULL, turnover.upper=10000, eps.upper=3, trans.upper=100,
ode.eps=0)

Arguments

Details

This function sets up and executes the original HiSSE model. The model closely follows diversitree, although here we employ modified optimization procedures. For example, rather than optimizing birth and death separately, hisse optimizes orthogonal transformations of these variables: we let tau = birth+death define "net turnover", and we let eps = death/birth define the “extinction fraction”. This reparameterization alleviates problems associated with overfitting when birth and death are highly correlated, but both matter in explaining the diversity pattern. As for data file format, hisse expects a two column matrix or data frame, with the first column containing the species names and the second containing the binary character information. Note that the order of the data file and the names in the “phylo” object need not be in the same order; hisse deals with this internally. Also, the character information should be binary and coded as 0 and 1, otherwise the function will misbehave. However, if the state for a species is unknown, a user can specify this with a 2, and the state will be considered maximally ambiguous.

To setup a model, users input vectors containing values to indicate how many free parameters are to be estimated for each of the variables in the model. For example, the “turnover.anc” input vector is set by default as c(1,1,0,0). This means for state 0 and state 1, we are allowing one free parameter to define the net turnover rate (birth+death) in the model. This is essentially a BiSSE model with fixed turnover rates. Now, say we want to include separate turnover rates for both states we would simply input c(1,2,0,0). The last two entries, which in the preceding example are set to zero, correspond to the hidden states; the third entry corresponds to a hidden state associated with observed state 0, such that 0A (hidden state absent) is the first entry, and 0B (hidden state present) is the third entry. So, to set up a model with three turnover rates, where we include a free parameter for a hidden state associated with state 0 we input c(1,2,3,0). A full model would thus be c(1,2,3,4), which corresponds to four separate net turnover rates, for states 0A (first entry), 1A (second entry), 0B (third entry), and 1B (fourth entry). Extinction fraction, or “eps.anc”, follows the same format, though including a zero for a state we want to include in the model corresponds to no extinction, which is the Yule equivalent. In general, we follow this format to make it easier to generate a large set of nested models. Once the model is specified, the parameters can be estimated using the subplex routine (default), or use a two-step process (i.e., sann=TRUE) that first employs a stochastic simulated annealing procedure, which is later refined using the subplex routine.

The “trans.rate” input is the transition model and has an entirely different setup than turnover and extinction rates. See TransMatMaker function for more details.

For user-specified “root.p”, you should specify the probability for each state. If you are doing a hidden model, there will be four states: 0A, 1A, 0B, 1B. So if you wanted to say the root had to be state 0, you would specify “root.p = c(0.5, 0, 0.5, 0)”.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Finally, the options “.beta” and “timeslice” are included, but neither have been tested – needless to say, use at your own risk (but really, though, you should probably forget that these options exist for the time being). The “.beta” provides a means for testing for time-varying rates, whereas “timeslice” splits the tree to allow the process to vary before and after some user defined time period. These options will be further developed in due course.

Value

hisse.old returns an object of class hisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Exports a likelihood function conditioned on the data and a named vector with the parameters for each of the models.

Usage

makeGeoHiSSELikelihood(phy, data, hidden.areas=0, f=c(1,1,1), assume.cladogenetic=TRUE, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, dt.threads=1, ode.eps = 0, 
bad.likelihood = exp(-300))

Arguments

Details

This function sets up and returns the likelihood for the GeoHiSSE model together with a vector of parameters. The likelihood function is conditioned on the observed data and will return a value of loglikelihood given a vector of parameter values. The length of the parameter vector as well as the order of the parameter vector cannot be changed. Please pay special attention to the length of the parameter vector and the names of the parameters provided by the “pars” element of the list.

Note that when the likelihood computation fails the function will return the value set as “bad.likelihood”.

Value

makeGeoHiSSELikelihood returns a list with elements:

Author(s)

Jeremy M. Beaulieu and Daniel S. Caetano

References

Caetano, D.S., B.C. O'Meara, and J.M. Beaulieu. 2018. Hidden state models improve state-dependent diversification approaches, including biogeographic models. Evolution, 72:2308-2324.

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Examples

library(diversitree)
library(hisse)
## Generate data:
pars <- c(0.1, 0.2, 0.03, 0.03, 0.01, 0.01)
set.seed(4) 
phy <- tree.bisse(pars, max.t=30, x0=0)
sim.dat <- data.frame(names(phy$tip.state), phy$tip.state)
## Get lik function:
lik.hisse <- makeHiSSELikelihood(phy = phy, data = sim.dat, hidden.states = FALSE)
likf <- lik.hisse$log.lik
pars <- lik.hisse$pars
## Set the parameter values. Note that we have turnover and eps, not speciation and extinction!
pars <- setNames(c(0.1+0.03,0.2+0.03,0.03/0.1,0.03/0.2,0.01,0.01), names(pars))
## Compute the log-likelihood for the model.
likf(pars)

Description

Exports a likelihood function conditioned on the data and a named vector with the parameters for each of the models.

Usage

makeHiSSELikelihood(phy, data, hidden.states=TRUE, null4=FALSE,
f=c(1,1), condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, ode.eps=0)

Arguments

Details

This function sets up and returns the likelihood for the HiSSE model together with a vector of parameters. The likelihood function is conditioned on the observed data and will return a value of loglikelihood given a vector of parameter values. The length of the parameter vector as well as the order of the parameter vector cannot be changed. The parameter values are provided in natural form but are log-transformed for the likelihood evaluation. Please pay special attention to the length of the parameter vector and the names of the parameters provided by the “pars” element of the list.

When the option “null4” is set to TRUE, then the likelihood returned is for the HiSSE null model with 4 hidden states. The returned list will include an additional element named “trans.mat.guide” which can be used as a reference for the meaning of the 32 transition parameters. Note that the original model in “hisse.null4” usually sets all these transitions to a single value. This helps model estimation since information is limited to estimate distinct transition rates between parameters.

Value

makeHiSSELikelihood returns a list with elements:

Author(s)

Jeremy M. Beaulieu and Daniel S. Caetano

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Examples

library(diversitree)
library(hisse)
## Generate data:
pars <- c(0.1, 0.2, 0.03, 0.03, 0.01, 0.01)
set.seed(4) 
phy <- tree.bisse(pars, max.t=30, x0=0)
sim.dat <- data.frame(names(phy$tip.state), phy$tip.state)
## Get lik function:
lik.hisse <- makeHiSSELikelihood(phy = phy, data = sim.dat, hidden.states = FALSE)
likf <- lik.hisse$log.lik
pars <- lik.hisse$pars
## Set the parameter values. Note that we have turnover and eps, not speciation and extinction!
pars <- setNames(c(0.1+0.03,0.2+0.03,0.03/0.1,0.03/0.2,0.01,0.01), names(pars))
## Compute the log-likelihood for the model.
likf(pars)

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginRecon.old(phy, data, f, pars, hidden.states=TRUE, four.state.null=FALSE, 
timeslice=NULL, condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, 
AIC=NULL, verbose=TRUE, n.cores=NULL)

Arguments

Details

Conducts ancestral state reconstruction on the original hisse.old and hisseNull4.old functions. In this implementation the marginal probability of state i for a focal node is simply the overall likelihood of the tree and data when the state of the focal node is fixed in state i. Note that the likeliest tip states can also be estimated: we observe state 1, but the underlying state could either be state 1A or 1B. Thus, for any given node or tip we traverse the entire tree as many times as there are states in the model. As the size of the tree grows, however, these repeated tree traversals can slow the calculation down considerably. For this reason, we allow the marginal calculation to be conducted in parallel across any number of independent computer processors.

For user-specified “root.p”, you should specify the probability for each state. If you are doing a hidden model, there will be four states: 0A, 1A, 0B, 1B. So if you wanted to say the root had to be state 0, you would specify “root.p = c(0.5, 0, 0.5, 0)”.

See help for “hisse.old” for more on other parameters for this function.

Value

MarginRecon returns an object of class hisse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginReconGeoSSE(phy, data, f, pars, hidden.states=1, 
assume.cladogenetic=TRUE, condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL, AIC=NULL, get.tips.only=FALSE,
verbose=TRUE, n.cores=NULL, dt.threads=1)

Arguments

Details

In this implementation the marginal probability of area i for a focal node is simply the overall likelihood of the tree and data when the area of the focal node is fixed in area i. Note that the likeliest tip areas can also be estimated: we observe area 11, but the underlying area could either be area 11A or 11B. Thus, for any given node or tip we traverse the entire tree as many times as there are areas in the model. As the size of the tree grows, however, these repeated tree traversals can slow the calculation down considerably. For this reason, we allow the marginal calculation to be conducted in parallel across any number of independent computer processors.

For user-specified “root.p”, you should specify the probability for each area. If you are doing a hidden model, the number of parameters will depend on the number of hidden states included. For a two classes model there are six areas: 00A, 11A, 01A, 00B, 11B, 01B. So if you wanted to say the root had to be area A, you would specify “root.p = c(0.5, 0, 0, 0.5, 0, 0)”. The root area is 00, but there is an equal chance for hidden states A or B.

See help for “GeoHiSSE” for more on other parameters for this function.

Value

MarginReconGeoSSE returns an object of class geohisse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Goldberg, E. E., L. T. Lancaster, and R. H. Ree. 2011. Phylogenetic Inference of Reciprocal Effects between Geographic Range Evolution and Diversification. Syst. Biol. 60:451-465.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginReconGeoSSE.old(phy, data, f, pars, hidden.areas=TRUE, 
assume.cladogenetic=TRUE, condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL, AIC=NULL, verbose=TRUE, 
n.cores=NULL)

Arguments

Details

In this implementation the marginal probability of area i for a focal node is simply the overall likelihood of the tree and data when the area of the focal node is fixed in area i. Note that the likeliest tip areas can also be estimated: we observe area 1, but the underlying area could either be area 1A or 1B. Thus, for any given node or tip we traverse the entire tree as many times as there are areas in the model. As the size of the tree grows, however, these repeated tree traversals can slow the calculation down considerably. For this reason, we allow the marginal calculation to be conducted in parallel across any number of independent computer processors.

For user-specified “root.p”, you should specify the probability for each area. If you are doing a hidden model, the number of parameters will depend on the number of hidden areas included. For a two classes model there are six areas: A0, B0, AB0, A1, B1, AB1. So if you wanted to say the root had to be area A, you would specify “root.p = c(0.5, 0, 0, 0.5, 0, 0)”. There is 'root area is A and there is an equal chance for hidden area A0 or A1'.

See help for “GeoHiSSE.old” for more on other parameters for this function.

Value

MarginReconGeoSSE.old returns an object of class geohisse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Goldberg, E. E., L. T. Lancaster, and R. H. Ree. 2011. Phylogenetic Inference of Reciprocal Effects between Geographic Range Evolution and Diversification. Syst. Biol. 60:451-465.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginReconHiSSE(phy, data, f, pars, hidden.states=1, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, 
includes.fossils = FALSE, k.samples = NULL, tip.fog=NULL, AIC=NULL, 
get.tips.only=FALSE, verbose=TRUE, n.cores=NULL, dt.threads=1)

Arguments

Details

This is the marginal reconstruction algorithm for the newer, faster version of HiSSE. In this implementation the marginal probability of state i for a focal node is simply the overall likelihood of the tree and data when the state of the focal node is fixed in state i. Note that the likeliest tip states can also be estimated: we observe state 1, but the underlying state could either be state 1A or 1B. Thus, for any given node or tip we traverse the entire tree as many times as there are states in the model. As the size of the tree grows, however, these repeated tree traversals can slow the calculation down considerably. For this reason, we allow the marginal calculation to be conducted in parallel across any number of independent computer processors.

See help for “hisse” for more on other parameters for this function.

Value

MarginReconHiSSE returns an object of class hisse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginReconMiSSE(phy, f, pars, hidden.states=1, fixed.eps=NULL,
condition.on.survival=TRUE,  root.type="madfitz", root.p=NULL, includes.fossils=FALSE, 
k.samples=NULL, strat.intervals=NULL, AIC=NULL, get.tips.only=FALSE, verbose=TRUE, 
n.cores=NULL, dt.threads=1)

Arguments

Details

In this implementation the marginal probability of state combination i for a focal node is simply the overall likelihood of the tree and data when the state of the focal node is fixed in state combination i.

See help for “MiSSE” for more on other parameters for this function.

Value

MarginReconMiSSE returns an object of class misse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Estimates the likeliest states for both internal nodes and tips of a phylogeny using the marginal reconstruction algorithm.

Usage

MarginReconMuHiSSE(phy, data, f, pars, hidden.states=1, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, includes.fossils = FALSE,
k.samples = NULL, AIC=NULL, get.tips.only=FALSE, verbose=TRUE, n.cores=NULL, dt.threads=1)

Arguments

Details

In this implementation the marginal probability of state combination i for a focal node is simply the overall likelihood of the tree and data when the state of the focal node is fixed in state combination i. Note that the likeliest tip states can also be estimated: we observe state 00, but the underlying state could either be 00A or 00B. Thus, for any given node or tip we traverse the entire tree as many times as there are states in the model. As the size of the tree grows, however, these repeated tree traversals can slow the calculation down considerably. For this reason, we allow the marginal calculation to be conducted in parallel across any number of independent computer processors.

For user-specified “root.p”, you should specify the probability for each state combination. If you are doing a hidden model, the number of parameters will depend on the number of hidden states included. For a two classes model there are eight states: 00A, 01A, 10A, 11A, 00B, 01B, 10B, and 11B. So if you wanted to say the root had to be in state 00, you would specify “root.p = c(0.5, 0, 0, 0, 0.5, 0, 0, 0, 0)”. There is 50 percent chance the root state is 00 and there is an equal chance for hidden state A or B.

See help for “MuHiSSE” for more on other parameters for this function.

Value

MarginRecon returns an object of class muhisse.states. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters' effect on speciation and extinction. Syst. Biol. 56:701-710.

Nakov, T., Beaulieu, J.M., and Alverson, A.J. 2019. Diatoms diversify and turn over faster in freshwater than marine environments. Evolution, doi: https://doi.org/10.1111/evo.13832.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Sets up and executes a MiSSE model (Missing State Speciation and Extinction) on a phylogeny.

Usage

MiSSE(phy, f=1, turnover=c(1,2), eps=c(1,2), fixed.eps=NULL, condition.on.survival=TRUE,
root.type="madfitz", root.p=NULL, includes.fossils=FALSE, k.samples=NULL, 
strat.intervals=NULL, sann=TRUE, sann.its=5000, sann.temp=5230, sann.seed=-100377, 
bounded.search=TRUE, max.tol=.Machine$double.eps^.50, starting.vals=NULL, 
turnover.upper=10000, eps.upper=3, trans.upper=100, restart.obj=NULL, ode.eps=0, 
dt.threads=1, expand.mode=FALSE)

Arguments

Details

One thing pointed out in the original HiSSE paper (Beaulieu & O'Meara, 2016) is that the trait-independent hisse model is basically a model for traits and a separate model for shifts in diversification parameters, much like BAMM (though without priors, discontinuous inheritance of extinction probability, or other mathematical foibles). The hidden states can drive different diversification processes, and the traits just evolve in a regular trait model. At that point, there is no harm in just dropping the trait (or analyzing separately) and just focusing on diversification driven by unknown factors. That is what this function does. It sets up and executes a completely trait-free version of a HiSSE model.

Thus, all that is required is a tree. The model allows up to 26 possible hidden states in diversification (denoted by A-Z). Transitions among hidden states are governed by a global transition rate, q. A "shift" in diversification denotes a lineage tracking some unobserved, hidden state. An interesting byproduct of this assumption is that distantly related clades can actually share the same discrete set of diversification parameters.

Note that "hidden state" is a shorthand. We do not mean that there is a single, discrete character that is solely driving diversification differences. There is some heritable "thing" that affects rates: but this could be a combination of body size, oxygen concentration, trophic level, and how many other species are competing in an area. This is true for HiSSE, but is especially important to grasp for MiSSE. It could be that there is some single discrete trait that drives everything; it's more likely that a whole range of factors play a role, and we just slice them up into discrete categories, the same way we slice up mammals into carnivore / omnivore / herbivore or plants into woody / herbaceous when the reality is more continuous.

As with hisse, we employ a modified optimization procedure. In other words, rather than optimizing birth and death separately, MiSSE optimizes orthogonal transformations of these variables: we let tau = birth+death define "net turnover", and we let eps = death/birth define the “extinction fraction”. This reparameterization alleviates problems associated with overfitting when birth and death are highly correlated, but both matter in explaining the diversity pattern.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Value

MiSSE returns an object of class misse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Executes a set of MiSSE models (Missing State Speciation and Extinction) on a phylogeny, varying the number of parameters for turnover and extinction fraction and stopping when models stop being very good.

Usage

MiSSEGreedy(phy, f=1, possible.combos = 
generateMiSSEGreedyCombinations(shuffle.start=TRUE), stop.deltaAICc=10, save.file=NULL, 
n.cores=NULL, chunk.size=10, check.fits=FALSE, remove.bad=FALSE, n.tries=2, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, includes.fossils=FALSE, 
k.samples=NULL, strat.intervals=NULL, sann=TRUE, sann.its=5000, sann.temp=5230, 
sann.seed=-100377,bounded.search=TRUE, max.tol=.Machine$double.eps^.50, 
starting.vals=NULL, turnover.upper=10000, eps.upper=3, trans.upper=100, restart.obj=NULL, 
ode.eps=0)

Arguments

Details

See the MiSSE function for description of the method overall. It requires a set number of hidden state categories, but finding the best number of categories can be hard. For example, one could 1 to 26 different turnover categories and 1 to 26 possible extinction fraction categories. For most cases, we suspect that it makes sense to have the number of extinction fraction categories either equal to the number of turnover categories or set to the same category over the tree, but there are actually a lot of possibilities: have turnover=c(1,2,3) and eps=c(1,2,1), for example, or turnover=c(1,1,1) and eps=c(1,2,3). This uses the generateMiSSEGreedyCombinations function to generate a very large set of these possible models, then runs them in increasing complexity. By default, it stops when the models stop getting being reasonable in AICc. This is NOT where the models stop being significant (if you're looking for significance, note you don't get that from AICc), but where they probably will not contribute much to the model averaged parameter estimates and so may not be worth the bother of searching further. However, there's no guarantee that this is a wise decision: you could stop with 10 turnover rates, and 11 and 12 are far worse for AICc, but it could be that 13 turnover rates are much better than anything else (for that matter, the best number of turnover rates could be 42, even though MiSSe's current code can only go up to 26). And of course, in reality, the truth is that there is an infinite set of hidden rate parameters: a passing cloud blocks a tiny bit of energy for photosynthesis, so for that moment the rate of extinction for plants underneath the shadow is a tiny bit higher. You should not be using this to test hypotheses about how many hidden factors there are, but rather as a way to get a good enough model to estimate rates on a tree.

You can change how quickly the function stops trying new models with stop.deltaAICc. Once it runs a chunk of models where the best gets a model that is at least stop.deltaAICc worse than the *current* best model, it stops running new models. Since this is based on current best AICc, and we start with simple models, there's an asymmetry: a terrible model with no rate variation is always included, but a slightly less terrible model with 26 turnover rates might never be run.

This works MUCH faster when run in parallel. To do so, set n.cores to the number of available cores on your machine (for example, for a modern Mac laptop, this would be 4). An easy way to do this automatically is to use n.cores=parallel::detectCores(). The default is one core; if running this function on a cluster, a default of parallel::detectCores() might take over an entire compute node when you're supposed to be using just one core, and get you in trouble. Since we run many models, the most natural approach is to run one model per core, see if at least one of the models are still ok, then send out the next models out to all the cores. Setting n.cores to the number of parallel jobs you want, and setting chunk.size to NULL, will do this. However, this is slightly inefficient – the odds are that some cores will finish earlier than others, and will be waiting until all finish. So a different approach is to set a chunk.size greater than n.cores – it will still use no more than n.cores at a time, but once one model in the set finishes it will send off the next until all models in the chunk are run. This keeps the computer even busier, but then it won't stop to check to make sure the models are still feasible as often, and it only saves intermediate results after each chunk of models finishes. Our recommendation is to use n.cores=parallel::detectCores() if you're on a machine where you can use all the cores. By default, we set chunk.size=10 so it looks at ten models before deciding none of them have a good enough AICc to keep looking at all models. If chunk.size is smaller, say, 2, it will look at only two models and if neither is within stop.deltaAICc of the best model, it will stop looking at the next chunk of models.

After every chunk of models are done, this function will display the status: what models have been run, what the likelihoods and AICs are, etc. It will also predict how long future runs will check (based on a linear regression between the number of free parameters and log(minutes to run)). These are just estimates based on the runs so far, but it's a stochastic search and can take more or less time.

Saving output while this goes is highly recommended – you can see how things are going and salvage something if a run fails (then, start again, deleting the models that worked from possible.combos and USING A DIFFERENT FILE TO SAVE TO SO YOU DON'T OVERWRITE THE OLD ONE. You can so this by giving a file name (including path, if you want) as the save.file argument. This will save the list of finished models (misse.list) and the possible.combos data.frame with additional information.

Value

MiSSEGreedy returns a list of class misse.fit objects.

Author(s)

Brian C. O'Meara

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Vasconcelos, T., B.C. O'Meara, and J.M. Beaulieu. 2022. A flexible method for estimating tip diversification rates across a range of speciation and extinction scenarios. Evolution, 76:1420-1433.

Description

A function for assessing whether models optimized well, identifying bad fits, and rerunning them if needed.

Usage

MiSSENet(misse.list, n.tries=2, remove.bad=TRUE, dont.rerun=FALSE, save.file=NULL, 
n.cores=1, sann=TRUE, sann.its=5000, sann.temp=5230, bounded.search=TRUE, 
starting.vals=NULL, turnover.upper=10000, eps.upper=3, trans.upper=100, restart.obj=NULL)

Arguments

Details

This function is used to triage poorly optimized models after a MiSSEGreedy run. It is normally invoked within MiSSEGreedy, but it can also be used as a standalone function, to simply identify poorly identify models and/or rerun them.

Author(s)

Jeremy M. Beaulieu

References

Vasconcelos, T., B.C. O'Meara, and J.M. Beaulieu. 2022. A flexible method for estimating tip diversification rates across a range of speciation and extinction scenarios. Evolution, 76:1420-1433.

Description

Summarizes reconstructions from a single model or a set of models and returns model averaged rates.

Usage

GetModelAveRates(x, AIC.weights=NULL, type=c("tips", "nodes", "both"),
bound.par.matrix=cbind(c(0,-1000,0,0,0),c(1000,1000,1000,3,1000)) )

Arguments

Details

Provides a data frame model-averaged rates for all possible configurations of the model parameters (i.e., turnover, net.div, speciation, extinction, or extinction.fraction), either for all tips or for all nodes. As with the plotting function, if you give a single hisse.state object, it uses that, and the rates account for uncertainty in the marginal probabilities of the states; if you give it a list of them, it will model-average the results (it assumes the trees are the same) in addition to accounting for uncertainty in the marginal probabilities at tips and nodes. If 'AIC.weights==NULL', then the models in the set do not require the '$AIC' element to compute the AICw. Otherwise the function will return an error message.

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

Sets up and executes a MuHiSSE model (Multicharacter Hidden State Speciation and Extinction) on a phylogeny and character distribution.

Usage

MuHiSSE(phy, data, f=c(1,1,1,1), turnover=c(1,2,3,4), eps=c(1,2,3,4), 
hidden.states=FALSE, trans.rate=NULL, condition.on.survival=TRUE, 
root.type="madfitz", root.p=NULL, includes.fossils=FALSE, k.samples=NULL, sann=TRUE, 
sann.its=1000, bounded.search=TRUE, max.tol=.Machine$double.eps^.50, starting.vals=NULL, 
turnover.upper=10000, eps.upper=3, trans.upper=100, restart.obj=NULL, ode.eps=0, 
dt.threads=1)

Arguments

Details

This function sets up and executes a multiple state HiSSE model. The model allows up to 8 hidden categories (hidden states A-H), and implements a more efficient means of carrying out the branch calculation. Specifically, we break up the tree into carry out all descendent branch calculations simultaneously, combine the probabilities based on their shared ancestry, then repeat for the next set of descendents. In testing, we've found that as the number of taxa increases, the calculation becomes much more efficient. In future versions, we will likely allow for multicore processing of these calculations to further improve speed. We also note that there is vignette that describes more details for running this particular function.

As for data file format, MuHiSSE expects a three column matrix or data frame, with the first column containing the species names and the second and third containing the binary character information. Note that the order of the data file and the names in the “phylo” object need not be in the same order; MuHiSSE deals with this internally. Also, the character information must be coded as 0 and 1, otherwise, the function will misbehave. However, if the state for a species is unknown for either character, a user can specify this with a 2, and the state will be considered maximally ambiguous for all relevant character combinations. For example, if character 1 is in state 0, but character 2 is provided a 2, then the program provides a probability of 1 for 00 and a probability of for 01.

As with hisse, we employ a modified optimization procedure. In other words, rather than optimizing birth and death separately, MuHisse optimizes orthogonal transformations of these variables: we let tau = birth+death define "net turnover", and we let eps = death/birth define the “extinction fraction”. This reparameterization alleviates problems associated with overfitting when birth and death are highly correlated, but both matter in explaining the diversity pattern.

To setup a model, users input vectors containing values to indicate how many free parameters are to be estimated for each of the variables in the model. This is done using the turnover and extinct.frac parameters. One needs to specify a value for each of the parameters of the model, when two parameters show the same value, then the parameters are set to be linked during the estimation of the model. For example, a MuHiSSE model with 1 hidden state and all free parameters has turnover = 1:8. The same model with turnover rates constrained to be the same for all hidden states has turnover = c(1,2,3,4,1,2,3,4). This same format applies to extinct.frac.

The “trans.rate” input is the transition model and has an entirely different setup than speciation and extinction rates. See TransMatMakerMuHiSSE function for more details.

For user-specified “root.p”, you should specify the probability for each state combination. If you are doing a hidden model, there will be eight state combinations: 00A, 01A, 10A, 11A, 00B, 01B, 10B, 11B. So if you wanted to say the root had to be in state 00, and since you do not know the hidden state, you would specify “root.p = c(0.5, 0, 0, 0, 0.5, 0, 0, 0)”. In other words, the root has a 50% chance to be in one of the states to be 00A or 00B.

For the “root.type” option, we are currently maintaining the previous default of “madfitz”. However, it was recently pointed out by Herrera-Alsina et al. (2018) that at the root, the individual likelihoods for each possible state should be conditioned prior to averaging the individual likelihoods across states. This can be set doing “herr_als”. It is unclear to us which is exactly correct, but it does seem that both “madfitz” and “herr_als” behave exactly as they should in the case of character-independent diversification (i.e., reduces to likelihood of tree + likelihood of trait model). We've also tested the behavior and the likelihood differences are very subtle and the parameter estimates in simulation are nearly indistinguishable from the “madfitz” conditioning scheme. We provide both options and encourage users to try both and let us know conditions in which the result vary dramatically under the two root implementations. We suspect they do not.

Also, note, that in the case of “root.type=user” and “root.type=equal” are no longer explicit “root.type” options. Instead, either “madfitz” or “herr_als” are specified and the “root.p” can be set to allow for custom root options.

Value

MuHiSSE returns an object of class muhisse.fit. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nakov, T., Beaulieu, J.M., and Alverson, A.J. 2019. Diatoms diversify and turn over faster in freshwater than marine environments. Evolution, doi: https://doi.org/10.1111/evo.13832.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

A plotting function for visualizing changes in states and rates over a phylogeny

Usage

## S3 method for class 'geohisse.states'
plot(x, rate.param = "net.div", type = "fan",
show.tip.label = TRUE, fsize = 1.0, legend = TRUE, ...)

Arguments

Details

Additional parameters can be defined using “...”:

“do.observed.only” is a logical indicating whether just the states should be plotted; for now, only TRUE works. “rate.colors” are user specified colors to be used for coloring rates. “state.colors” are user specified colors to be used for coloring states. A vector with the color for state 1 (endemic area '0'), state 2 (endemic area '1') and state 0 (widespread area '01'). Always in this order! “edge.width” is the width of the branches of the phylogeny. “width.factor” is the factor multiplying the “edge.width” to get the width of the branch for the states tree. Needs to be a numeric value between 0 and 1. “rate.range” is an optional two element vector. If present, specifies the range of rate values to use for plotting. “lims.percentage.correction” deals with cases where the limits are slightly smaller than the values due to imprecision issues. “legend.position” are the coordinates for placing the legend. “legend.cex” is the text size inside the legend. “legend.kernel” lets you specify the way the density plot or histogram is made for rates. A value of “auto” chooses what we think is the best option given your data, “hist” makes a histogram, “rectangular”, “gaussian”, and others make a density plot. See ?density for all non-“hist” options. “legend.bg” sets the color for the legend background. “mar” sets the margins for the plot. See more information in 'help(par)'. “outline” is whether to plot an outline on the branches of the tree. Choose the color of the outline using the parameter outline.color. “outline.color” is the color for the outline. Defaults to "black". “swap.underscore” sets whether to substitute all "_" with " " on the labels for the tips.

This function is very similar to the hisse::plot.hisse.states function. See more details in help page for hisse::plot.hisse.states function.

Author(s)

Brian O'Meara and Daniel Caetano

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

A plotting function for visualizing changes in states and rates over a phylogeny

Usage

## S3 method for class 'hisse.states'
plot(x, rate.param = "net.div", type = "fan",
show.tip.label = TRUE, fsize = 1.0, legend = "tips", ...)

Arguments

Details

Provides phylogeny that shows a heat map of the diversification rate parameter you specify (which could be turnover, net.div, speciation, extinction, or extinction.fraction). The discrete state reconstruction appears as lines on top of the heat map. If you give a single hisse.state object, it uses that; if you give it a list of them, it will model-average the results (it assumes the trees are the same). Colors can be specified by sending a vector of colors to rate.colors or state.colors (the defaults are red to blue for rate and white to black for state). You can specify two or more colors: c("red", "gray", "blue") for example. By default the visualization uses the minimum rate on the tree for the minimum color, and the maximum rate for the maximum color, but you may want to use the same color scale across models, even if some of them have a smaller range than others. To do this, pass a vector with the minimum and maximum rate across all models to the visualization for all models and they will use the same scale. There are many options for adding a legend. A traditional legend showing what values a color corresponds to is “traditional”, like what plotSimmap will show in phytools. However, we can also use the legend to show the distribution of values, rather than just a key to color. “tips” shows a density plot of states or rates at tips, “internal” a distribution at internal nodes, and “all” at all nodes in the tree. For the density or histogram plots, you can let the package pick the best visualization or choose yourself whether to use a histogram or density plot, and if the latter, what kernel you want. The legend can be moved around the overall tree plot by using “legend.position”: this is a vector that specifies the “fig” argument to “par”: c(x1, x2, y1, y2), where the values are the starting and ending positions as a fraction of the overall plot. By default, the legend starts at the lower left corner and continues up 20 the rest of the plot (c(0, 0.2, 0, 0.2)): by changing values, you can make the legend larger or smaller and change its position. The heatmap code is modified slightly from Liam Revell's phytools package.

Additional parameters to control the plot: “do.observed.only” is a logical indicating whether just the states should be plotted; for now, only TRUE works. “rate.colors” sets user specified colors to be used for coloring rates. “state.colors” sets user specified colors to be used for coloring states. “edge.width” sets the width of the branches of the phylogeny. “width.factor” is the factor multiplying the edge.width to get the width of the branch for the states tree. Needs to be a numeric value between 0 and 1. “rate.range” is an optional two element vector. If present, specifies the range of rate values to use for plotting. “lims.percentage.correction” deals with cases where the limits are slightly smaller than the values due to imprecision issues. “legend.position” are the coordinates for placing the legend. “legend.cex” is the text size inside the legend. “legend.kernel.rates” used for legend=tips, internal, or all, lets you specify the way the density plot or histogram is made for rates. “auto” chooses what we think is the best option given your data, “hist” makes a histogram, “rectangular”, “gaussian”, and others make a density plot. See ?density for all non-“hist” options. “legend.kernel.states” is as above, for states. “legend.bg” sets the color for the legend background. “mar” sets the margins for the plot. See more information in 'help(par)'. “outline” sets whether to plot an outline on the branches of the tree. Choose the color of the outline using the parameter outline.color. “outline.color” sets the color for the outline. Defaults to "black". “swap.underscore” defines whether to substitute all "_" with " " on the labels for the tips.

Author(s)

Brian O'Meara and Daniel Caetano

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

A plotting function for visualizing changes in states and rates over a phylogeny

Usage

## S3 method for class 'misse.states'
plot(x, rate.param = "net.div", type = "fan",
show.tip.label = TRUE, fsize = 1.0, legend = "tips", ...)

Arguments

Details

Additional parameters can be defined using “...”:

“do.observed.only” is a logical indicating whether just the states should be plotted; for now, only TRUE works. “rate.colors” are user specified colors to be used for coloring rates. “state.colors” are user specified colors to be used for coloring states. A vector with the color for states 00, 01, 10, and 11 (in this order). “edge.width” is the width of the branches of the phylogeny. “width.factor” is the factor multiplying the “edge.width” to get the width of the branch for the states tree. Needs to be a numeric value between 0 and 1. “rate.range” is an optional two element vector. If present, specifies the range of rate values to use for plotting. “lims.percentage.correction” deals with cases where the limits are slightly smaller than the values due to imprecision issues. “legend.position” are the coordinates for placing the legend. “legend.cex” is the text size inside the legend. “legend.kernel” lets you specify the way the density plot or histogram is made for rates. A value of “auto” chooses what we think is the best option given your data, “hist” makes a histogram, “rectangular”, “gaussian”, and others make a density plot. See ?density for all non-“hist” options. “legend.bg” sets the color for the legend background. “mar” sets the margins for the plot. See more information in 'help(par)'. “outline” is whether to plot an outline on the branches of the tree. Choose the color of the outline using the parameter outline.color. “outline.color” is the color for the outline. Defaults to "black". “swap.underscore” sets whether to substitute all "_" with " " on the labels for the tips.

This function is very similar to the hisse::plot.hisse.states function. See more details in help page for hisse::plot.hisse.states function.

Author(s)

Brian O'Meara and Daniel Caetano

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

A plotting function for visualizing changes in states and rates over a phylogeny

Usage

## S3 method for class 'muhisse.states'
plot(x, rate.param = "net.div", type = "fan",
show.tip.label = TRUE, fsize = 1.0, legend = TRUE, ...)

Arguments

Details

Additional parameters can be defined using “...”:

“do.observed.only” is a logical indicating whether just the states should be plotted; for now, only TRUE works. “rate.colors” are user specified colors to be used for coloring rates. “state.colors” are user specified colors to be used for coloring states. A vector with the color for states 00, 01, 10, and 11 (in this order). “edge.width” is the width of the branches of the phylogeny. “width.factor” is the factor multiplying the “edge.width” to get the width of the branch for the states tree. Needs to be a numeric value between 0 and 1. “rate.range” is an optional two element vector. If present, specifies the range of rate values to use for plotting. “lims.percentage.correction” deals with cases where the limits are slightly smaller than the values due to imprecision issues. “legend.position” are the coordinates for placing the legend. “legend.cex” is the text size inside the legend. “legend.kernel” lets you specify the way the density plot or histogram is made for rates. A value of “auto” chooses what we think is the best option given your data, “hist” makes a histogram, “rectangular”, “gaussian”, and others make a density plot. See ?density for all non-“hist” options. “legend.bg” sets the color for the legend background. “mar” sets the margins for the plot. See more information in 'help(par)'. “outline” is whether to plot an outline on the branches of the tree. Choose the color of the outline using the parameter outline.color. “outline.color” is the color for the outline. Defaults to "black". “swap.underscore” sets whether to substitute all "_" with " " on the labels for the tips.

This function is very similar to the hisse::plot.hisse.states function. See more details in help page for hisse::plot.hisse.states function.

Author(s)

Brian O'Meara and Daniel Caetano

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Description

A plotting function for visualizing the model space explored in MiSSEGreedy. Arguments in ... are passed to plot.igraph.

Usage

PlotMisseSpace(x, possible.combos=NULL, arrows.by.weight=FALSE, ...)

Arguments

Details

If the input x is a list of misse fits from MiSSEGreedy then size of the vertices are in proportion of their AIC weights. If only the “possible.combos” is input, and x=NULL, then the model space will be plotted but with each vertex having the same size.

Author(s)

Jeremy M. Beaulieu

References

Vasconcelos, T., B.C. O'Meara, and J.M. Beaulieu. 2022. A flexible method for estimating tip diversification rates across a range of speciation and extinction scenarios. Evolution, 76:1420-1433.

Description

Takes output from GetFossils and places extinct lineages in the tree and produces a properly formatted table of fossil samples from lineages that survive to the present (k fossils).

Usage

ProcessSimSample(phy, f)

Arguments

Value

Returns an object that contains:

Description

Takes output from GetFossils and places extinct lineages in the tree and produces a properly formatted table of stratigraphic intervals.

Usage

ProcessSimStrat(phy, f)

Arguments

Value

Returns an object that contains:

Description

Removes models that fit equivalently well to models that are as simple or simpler.

Usage

PruneRedundantModels(..., precision=1e-5)

Arguments

Details

Imagine two models: Model 1: parameters a and b allowed to vary Model 2: parameters a set to equal b

If the maximum likelihood estimate for a and b in model 1 are that they have the same value, it's essentially the same as including model 2 twice, and so other models have lower weight in consequence. This comes up with hidden state models where some hidden states are in extremely low frequencies throughout: they have more parameters than simpler models but the likelihood is essentially the same. Burnham and Anderson (2003, page 342ff) recommend removing redundant models in cases like this where two models are effectively identical.

This function does this by ordering models by the number of free parameters, then deleting models that have log likelihoods nearly exactly equal (to the precision set above) to a model that is as simple or simpler. This is not a perfect solution – it's possible that models could have very different parameters and have the same likelihood, for example. Also the "as simple" means that if there are two models with equal numbers of parameters, whichever is first gets saved and the second is removed.

Author(s)

Brian O'Meara

Description

Converts the $results element of the return to a phylo object

Usage

SimToPhylo(results, include.extinct=FALSE, drop.stem=TRUE)

Arguments

Details

This takes the $results object from a SimulateHisse run and converts it to an ape phylo object. If there are no taxa on the final tree, it returns NA; if there is one taxon, it returns a one taxon tree. This is behavior different from diversitree's tree simulators, which returns NULL in both the zero and one taxon case. Extinct taxa can be pruned or not. For simulations starting with one taxon, and/or for simulations with some extinction, the final tree can have a time when there is a single lineage before it radiates into the crown group. This stem can be included or not. The tip states are stored in $tip.state in the returned tree.

Value

Author(s)

Brian O'Meara

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Examples

simulated.result <- SimulateHisse(c(.3, .1), c(.1, 0), 
matrix(c(NA, 0.2, .3, NA), nrow=2), max.taxa=35, x0=1)	
par(mfcol=c(1,2))
plot(SimToPhylo(simulated.result$results, include.extinct=TRUE))
plot(SimToPhylo(simulated.result$results, include.extinct=FALSE))

Description

Simulation function for the GeoHiSSE model

Usage

SimulateGeoHiSSE(pars, hidden.traits=1, x0="0A", max.taxa=Inf,
max.time=Inf, add.jumps=FALSE, add.extinction=FALSE, 
include.extinct=FALSE, return.GeoHiSSE_pars=FALSE, 
override.safeties=FALSE)

Arguments

Details

This function uses diversitree::tree.classe function to simulate the tree under the GeoSSE model with hidden states. The parameter pars is a list of class "GeoHiSSE_pars" that can be generated using this same function. The best way is to first run SimulateGeoHiSSE with empty parameters and return.GeoHiSSE_pars=TRUE and with hidden.areas equal to the number of desired hidden states in the model. This will return an object in list format with a set of four matrices. Each matrix controls a different set of parameters of the model. All parameters are set to 0 and parameters not used by the model are set to NA. To prepare the simulation, edit these matrices and change the desired parameter values. Leaving a parameter equal to 0 is the same as reducing the model by eliminating this parameter of the model.

Value

This returns a list, with the following elements:

Author(s)

Daniel Caetano

Examples

## Not run: 
## Get the a list with the correct parameters to run the simulation:
pars <- SimulateGeoHiSSE(hidden.areas=1, return.GeoHiSSE_pars=TRUE)
## Edit the parameter values:
pars$model.pars[1:3,] <- 0.1
pars$model.pars[4:5,] <- 0.05
pars$model.pars[6:7,] <- 0.01
pars$q.01[1,2] <- pars$q.01[2,1] <- 0.01
pars$q.1[1,2] <- pars$q.1[2,1] <- 0.02
pars$q.2[1,2] <- pars$q.2[2,1] <- 0.02
## Run the simulation:
sim <- SimulateGeoHiSSE(pars=pars, hidden.areas=1, x0="01A", max.taxa=100)

## End(Not run)

Description

Flexible simulation function allowing checkpointing

Usage

SimulateHisse(turnover.rates, eps.values, transition.rates, max.taxa=Inf, max.t=Inf, 
max.wall.time=Inf, x0, nstart=1, checkpoint.file=NULL, checkpoint.frequency=100, 
checkpoint.start.object=NULL, override.safeties=FALSE, mass.extinction.heights=c(),
mass.extinction.magnitudes=c())

Arguments

Details

Note that currently, the simulator assumes turnover.rates, eps.values, and transition rates are in the same state order. Remember that turnover.rates are birth + death rates, and eps.values are death / birth rates.

We strongly advise putting some limits to stop the run. These can be any combination of max.taxa, max.t, and max.wall.time. For example, you could set up the run to stop when you hit 1000 taxa or 3600 seconds (one hour) of running on your computer, whichever comes first. If you want to run with no limits, you have to specify override.safeties=TRUE, and you should know what you're doing here (the runs might never finish). There is only one starting state (x0), which should be the index into the rate vectors, starting with 0: that is, if your states are 0A, 0B, 1A, and 1B, x0=0 would start in 0A, x0=3 would start in 1B. nstart lets you choose how many taxa to start with: it could be one lineage to start from a single taxon, in which case you'll have to wait some time for the first speciation event, or you could start with two, to start simulation with a clade of this size (though, if extinction is nonzero, you are not guaranteed to have the final crown group include both of these descendants). You can choose numbers higher than two, to start with, say, a 50-species polytomy. We're not sure why you would. You can add checkpointing: runs take a long time, and if it crashes, this will let you start from the same point in the last saved file (though with a different seed, unless you specify this yourself). If you want to save checkpoints as you go along, specify a file in checkpoint.file. Every checkpoint.frequency events, where an event is a speciation, extinction, or transition event, it will save current progress to a file. The smaller this value, the more frequently the simulation will be saved: fewer lost steps if you have to re-simulate, but slower during the run because it spends time writing to disk. If you want to start from a checkpointed analysis, load the file into R and then specify the object within R [not the file] containing the checkpointed results (by default, saved in an object called checkpoint.result) and the simulation will continue from that point.

Results are returned in a list, containing a dataframe with the outcome of the simulation (one row per edge in the tree, and containing information on the lengths, ancestor, tipward height from the original root, and state at the tipward end of the edge (you can get state at the rootward end by getting the tipward state of the ancestral edge). It may be of interest to know how many events of each type occurred, which is saved in $birth.counts, $death.counts, $transition.counts. The total number of surviving taxa is stored in $n.surviving.

The output can be returned as an ape phylo tree by passing the $results element of the final output to SimToPhylo().

Value

This returns a list, with the following elements:

Author(s)

Brian O'Meara

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

Examples

simulated.result <- SimulateHisse(c(.3, .1), c(.1, 0), 
matrix(c(NA, 0.2, .3, NA), nrow=2), max.taxa=35, x0=1)	
par(mfcol=c(1,2))
plot(SimToPhylo(simulated.result$results, include.extinct=TRUE))
plot(SimToPhylo(simulated.result$results, include.extinct=FALSE))

Description

This provides an overview of all the examined models, including (optionally) a reconstruction of the diversification parameters across the models.

Usage

SummarizeMiSSEGreedy(greedy.result, min.weight=0.01, n.cores=1, recon=TRUE)

Arguments

Details

After doing a MiSSEGreedy run, this function provides an overview of the models. The overview object is a data.frame with columns for the model number, its AICc and related measures, the number of free parameters, how long that model took, and whether it is used to reconstruct the diversification parameters. Whether to include a model or not in a reconstruction is up to the user: including very bad models can lead to a reconstruction that is not very good (the model might have very low weight, but if the parameter estimate is still near infinity, for example, it could have a major impact). By default we use models with a weight of at least 0.01.

The rates object is a 3d array: the dimensions are the tips and internal nodes, the parameters being estimated, and the model(s) being used. For example, if you store the SummarizeMiSSEGreedy() output as summarized_results, then summarized_results$rates[,"extinction.fraction","best"] is the extinction fraction estimates for the tips and internal nodes from the best model.

Value

SummarizeMiSSEGreedy returns a list of containing a data.frame with an overview of models (overview) and an array with rates (rates).

Author(s)

Brian C. O'Meara

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Herrera-Alsina, L., P. van Els, and R.S. Etienne. 2018. Detecting the dependence of diversification on multiples traits from phylogenetic trees and trait data. Systematic Biology, 68:317-328.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegion.old(hisse.old.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, output.type="turnover", hidden.states=TRUE, 
condition.on.survival=TRUE, root.type="madfitz", root.p=NULL, verbose=TRUE)

Arguments

Details

This function provides a means for sampling the likelihood surface quickly to estimate confidence intervals that reflect the uncertainty in the MLE. The function starts with the MLE from the hisse run. It then uses a scaling multiplier to generate an interval by which to randomly alter each parameter. However, the algorithm was designed to “feel” the boundaries of the random search. In other words, when the algorithm begins to sample the hinterlands of the surface, it will know to restrict the boundary to allow sampling of more reasonable values based on the currently sampled set. The goal of this sampling process is to find points within some desired distance from the MLE; by default we assume this distance is 2 lnLik. The confidence interval can be estimated directly from these points. The full set of points tried are also provided and can be used to generate contour plots (though, it is not entirely straightforward to do so – but certainly doable).

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegion.old returns an object of class hisse.old.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegionGeoSSE(geohisse.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, verbose=TRUE)

Arguments

Details

This function provides a means for sampling the likelihood surface quickly to estimate confidence intervals that reflect the uncertainty in the MLE. The function starts with the MLE from the hisse run. It then uses a scaling multiplier to generate an interval by which to randomly alter each parameter. However, the algorithm was designed to “feel” the boundaries of the random search. In other words, when the algorithm begins to sample the hinterlands of the surface, it will know to restrict the boundary to allow sampling of more reasonable values based on the currently sampled set. The goal of this sampling process is to find points within some desired distance from the MLE; by default we assume this distance is 2 lnLik. The confidence interval can be estimated directly from these points. The full set of points tried are also provided and can be used to generate contour plots (though, it is not entirely straightforward to do so – but certainly doable).

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegionGeoSSE.old returns an object of class geohisse.old.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegionGeoSSE.old(geohisse.old.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, verbose=TRUE)

Arguments

Details

This function provides a means for sampling the likelihood surface quickly to estimate confidence intervals that reflect the uncertainty in the MLE. The function starts with the MLE from the hisse run. It then uses a scaling multiplier to generate an interval by which to randomly alter each parameter. However, the algorithm was designed to “feel” the boundaries of the random search. In other words, when the algorithm begins to sample the hinterlands of the surface, it will know to restrict the boundary to allow sampling of more reasonable values based on the currently sampled set. The goal of this sampling process is to find points within some desired distance from the MLE; by default we assume this distance is 2 lnLik. The confidence interval can be estimated directly from these points. The full set of points tried are also provided and can be used to generate contour plots (though, it is not entirely straightforward to do so – but certainly doable).

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegionGeoSSE.old returns an object of class geohisse.old.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegionHiSSE(hisse.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, verbose=TRUE)

Arguments

Details

This is the support region estimator for the new version of hisse.

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegionHiSSE returns an object of class hisse.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegionMiSSE(misse.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, verbose=TRUE)

Arguments

Details

This function provides a means for sampling the likelihood surface quickly to estimate confidence intervals that reflect the uncertainty in the MLE. The function starts with the MLE from the hisse run. It then uses a scaling multiplier to generate an interval by which to randomly alter each parameter. However, the algorithm was designed to “feel” the boundaries of the random search. In other words, when the algorithm begins to sample the hinterlands of the surface, it will know to restrict the boundary to allow sampling of more reasonable values based on the currently sampled set. The goal of this sampling process is to find points within some desired distance from the MLE; by default we assume this distance is 2 lnLik. The confidence interval can be estimated directly from these points. The full set of points tried are also provided and can be used to generate contour plots (though, it is not entirely straightforward to do so – but certainly doable).

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegionMiSSE returns an object of class misse.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Adaptively samples points for each parameter to obtain an estimate of the confidence intervals.

Usage

SupportRegionMuHiSSE(muhisse.obj, n.points=1000, scale.int=0.1, desired.delta=2, 
min.number.points=10, verbose=TRUE)

Arguments

Details

This function provides a means for sampling the likelihood surface quickly to estimate confidence intervals that reflect the uncertainty in the MLE. The function starts with the MLE from the hisse run. It then uses a scaling multiplier to generate an interval by which to randomly alter each parameter. However, the algorithm was designed to “feel” the boundaries of the random search. In other words, when the algorithm begins to sample the hinterlands of the surface, it will know to restrict the boundary to allow sampling of more reasonable values based on the currently sampled set. The goal of this sampling process is to find points within some desired distance from the MLE; by default we assume this distance is 2 lnLik. The confidence interval can be estimated directly from these points. The full set of points tried are also provided and can be used to generate contour plots (though, it is not entirely straightforward to do so – but certainly doable).

Note the scale.int option. This roughly sets the variance for sampling points. If this seems to take a long while to find enough points within the desired likelihood region consider reducing scale.int to either 0.05 or, in some cases, 0.01.

Value

SupportRegionMuHiSSE returns an object of class muhisse.support. This is a list with elements:

Author(s)

Jeremy M. Beaulieu

References

Beaulieu, J.M, and B.C. O'Meara. 2016. Detecting hidden diversification shifts in models of trait-dependent speciation and extinction. Syst. Biol. 65:583-601.

FitzJohn R.G., Maddison W.P., and Otto S.P. 2009. Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies. Syst. Biol. 58:595-611.

Maddison W.P., Midford P.E., and Otto S.P. 2007. Estimating a binary characters effect on speciation and extinction. Syst. Biol. 56:701-710.

Nakov, T., Beaulieu, J.M., and Alverson, A.J. 2019. Diatoms diversify and turn over faster in freshwater than marine environments. Evolution, doi: https://doi.org/10.1111/evo.13832.

Nee S., May R.M., and Harvey P.H. 1994. The reconstructed evolutionary process. Philos. Trans. R. Soc. Lond. B Biol. Sci. 344:305-311.

Description

Performs linear regression between phylogenetic independent contrasts (PICs) of tip rates and continuous trais

Usage

TipCorrelation(phy, tip.rate, trait, log=TRUE, remove.cherries=TRUE, 
scaled=TRUE, positivise=TRUE, use.lmorigin=TRUE)

Arguments

Details

Tip rates as those obtained with MiSSEGreedy are analogous to a continuous trait evolving in the tree and must also be corrected for phylogenetic non-independence before regression analyses. This function does that by using PICs (Felsenstein, 1985) and gives the additional option of removing PICs from 'cherries' from analyses. Cherries (theoretically) inherit the exact same rate class probabilities in any model that uses just branch lengths to estimate tip rates. For that reason, they may: (1) present identical tip rates, forcing the slope of the regression to be close to 0 since all PICs for cherries will be 0; and (2) constitute pseudoreplicates in the analyses. We suspect that, for that reason, it may make sense to prune them out from any PIC analyses that uses tip-rate correlations from tree-only diversification approaches (see Vasconcelos et al. in prep.).

Value

Returns an object that contains:

Author(s)

Thais Vasconcelos and Brian O'Meara

References

Vasconcelos, T, B.C. O'Meara, and J.M. Beaulieu. In prep. Felsenstein, J. (1985). Phylogenies and the comparative method. The American Naturalist, 125, 1-15.

Description

Simulate key of parnames to translate between GeoHiSSE and ClaSSE models

Usage

TranslateParsMakerGeoHiSSE(k, add.extinction, add.jumps)

Arguments

Details

Function will only return the parameters that are relevant for a GeoHiSSE model. The ClaSSE model will have many more parameters. The extra parameters of the ClaSSE model need to be all set to 0. The number of parameters of the correspondent ClaSSE model given the parameter k is given by:

(k+1)*3

Value

Function returns a matrix of characters names. Column 'classe.pars' have the parameter names for a ClaSSE model and column 'geohisse.pars' have the parameter names for the matching GeoHiSSE model.

Author(s)

Daniel Caetano

Description

Generates and manipulates the index of the rate parameters to be optimized

Usage

TransMatMaker.old(hidden.states=FALSE)
ParDrop(rate.mat.index=NULL, drop.par=NULL)
ParEqual(rate.mat.index=NULL, eq.par=NULL)

Arguments

Details

Outputs the full index of the rate parameters that are to be optimized. The intention is that a user might want to see how the matrix is designed prior to an analysis and perhaps drop a few parameters beforehand due to some hypothesis that he or she might have. The resulting matrix is to be plugged directly into hisse.old.

Value

Returns a rate matrix index

Description

Generates and manipulates the index of the transition rate parameters to be optimized.

Usage

TransMatMakerGeoHiSSE(hidden.traits=0, make.null=FALSE, include.jumps=FALSE, 
	separate.extirpation=FALSE)

Arguments

Details

Outputs the full index of the rate parameters that are to be optimized.

The intention is that a user might want to see how the matrix is designed prior to an analysis and perhaps drop a few parameters beforehand due to some hypothesis that she might have. The resulting matrix is to be plugged directly into GeoHiSSE.

Value

Returns a rate matrix index.

Description

Generates and manipulates the index of the transition rate parameters to be optimized.

Usage

TransMatMakerHiSSE(hidden.traits=0, make.null=FALSE, cat.trans.vary=FALSE)

Arguments

Details

Outputs the full index of the rate parameters that are to be optimized.

The intention is that a user might want to see how the matrix is designed prior to an analysis and perhaps drop a few parameters beforehand due to some hypothesis that she might have. The resulting matrix is to be plugged directly into hisse.

Value

Returns a rate matrix index.

Description

Generates and manipulates the index of the transition rate parameters to be optimized.

Usage

TransMatMakerMuHiSSE(hidden.traits=0, make.null=FALSE, include.diagonals=FALSE, 
cat.trans.vary=FALSE)

Arguments

Details

Outputs the full index of the rate parameters that are to be optimized.

The intention is that a user might want to see how the matrix is designed prior to an analysis and perhaps drop a few parameters beforehand due to some hypothesis that she might have. The resulting matrix is to be plugged directly into MuHiSSE.

Value

Returns a rate matrix index.