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Clark, JS, C Nunes, and B Tomasek. 2019. Masting as an unreliable resource: spatio-temporal host diversity merged with consumer movement, storage, and diet. Ecological Monographs, e01381.
Qiu, T. …, and J. S. Clark. 2023. Mutualist dispersers and the global distribution of masting: mediation by climate and fertility. Nature Plants 9, 1044–1056.
mastif
uses seed traps and crop counts on trees to
estimate seed productivity and dispersion. Attributes of individual
trees and their local environments could explain their differences in
fecundity. Inference requires information on locations of trees and seed
traps and predictors ( covariates and factors ) that could explain
source strength.
Many trees are not reproductively mature, and reproductive status is often unknown. For individuals of unknown maturation status, the maturation status must be estimated together with fecundity.
Predictors of fecundity include individual traits, site characteristics, climate, synchronicity with other trees, and lag effects. For studies that involve multiple plots and years, there can be effects of climate between sites and through time. There may be synchronicity between individuals both within and between plots.
Observations incorporate two types of redistribution. The first type is a redistribution from species to seed types. Not all seed types can be definitely assigned to species. The contribution of trees of each species to multiple seed types must be estimated.
The second redistribution from source trees to seed traps. Within a plot-year there is dependence between all seed traps induced by a redistribution kernel, which describes how seeds produced by trees are dispersed in space. Because ‘seed shadows’ of trees overlap, seeds are assigned to trees only in a probabilistic sense. The capacity to obtain useful estimates depends on the number of trees, the number of seed traps, the dispersal distances of seeds, the taxonomic specificity of seed identification, and the number of species contributing to a given seed type.
In addition to covariates and factors, the process model for fecundity can involve random tree effects, random group effects ( e.g., species-location ), year effects, and autoregressive lags limited only by the duration of data. Dispersal estimtes can include species and site differences ( Clark et al. 2013 ).
mastif
simulates the posterior distributions of
parameters and latent variables using Gibbs sampling, with direct
sampling, Hamiltonian Markov chain ( HMC ) updates, and Metropolis
updates. mastif
is implemented in R and C++ with
Rcpp
and RcppArmadillo
libraries.
mastif
inputsThe model requires a minimum of six inputs directly as arguments to
the function mastif
. There are two formula
s,
two character vector
s, and four
data.frame
s:
Table 2. Basic inputs
object | mode |
components |
---|---|---|
formulaFec |
formula |
fecundity model |
formulaRep |
formula |
maturation model |
specNames |
character vector |
names in column species of
treeData to include in model |
seedNames |
character vector |
names of seed types to include in model: match column
names in seedData |
treeData |
data.frame |
tree-year by variables, includes columns
plot , tree , year ,
diam , repr |
seedData |
data.frame |
trap-year by seed types, includes columns
plot , trap , year , one for each
specNames |
xytree |
data.frame |
tree by location, includes columns plot ,
tree , x , y |
xytrap |
data.frame |
trap by location, includes columns plot ,
trap , x , y |
I introduce additional inputs in the sections that follow, but start with this basic model.
Data simulation is recommended as the place to start. A simulator provides insight on how well parameters from be identified from my data. The inputs to the simulator can specify the following:
Table 3. Simulation inputs to
mastSim
mastSim input |
explanation |
---|---|
nplot |
number of plots |
nyr |
mean no. years per plot |
ntree |
mean no. trees per plot |
ntrap |
mean no. traps per plot |
specNames |
tree species codes used in treeData
column |
seedNames |
seed type codes: a column name in
seedData |
Most of these inputs are stochasiticized to vary structure of each
data set. Here are inputs to the simulator for a species I’ll call
acerRubr
:
seedNames <- specNames <- 'acerRubr'
sim <- list( nplot=5, nyr=10, ntree=30, ntrap=40, specNames = specNames,
seedNames = seedNames )
The list sim
holds objects needed for simulation. Here
is the simulation:
inputs <- mastSim( sim ) # simulate dispersal data
seedData <- inputs$seedData # year, plot, trap, seed counts
treeData <- inputs$treeData # year, plot, tree data
xytree <- inputs$xytree # tree locations
xytrap <- inputs$xytrap # trap locations
formulaFec <- inputs$formulaFec # fecundity model
formulaRep <- inputs$formulaRep # maturation model
trueValues <- inputs$trueValues # true states and parameter values
I first summarize these model inputs
generated by
mastSim
.
mastSim
mastSim
generates inputs needed for model fitting with
mast
. These objects are simulated data, including the four
data.frame
s ( treeData
, seedData
,
xytree
, xytrap
) and formulas (
formulaFec
, formulaRep
). Other objects are
“true” parameter values and latent states in the
list truevalues
used to simulate the data (
fec
, repr
, betaFec
,
betaRep
, upar
, R
). I want to see
if mast
can recover these parameter values for the type of
data I simulated.
Here is a mapping of objects created by mastSim
to the
model in Clark et al. ( 2019 ):
Table 4. Some of the objects from the
list mastSim
.
mastSim object |
variable | explanation |
---|---|---|
trueValues$fec |
\(\psi_{ij, t}\) | conditional fecundity |
trueValues$repr |
\(\rho_{ij, t}\) | true maturation status |
treeData$repr |
\(z_{ij, t}\) | observed maturation status ( with NA
) |
trueValues$betaFec |
\(\beta^x\) | coefficients for fecundity |
trueValues$betaRep |
\(\beta^v\) | coefficients for maturation |
trueValues$R |
\(\mathbf{m}\) | specNames to
seedNames matrix , rows = \(\mathbf{m}_h\) |
seedData$active |
in \(A_{sj, t}\) | fraction of time trap is active |
seedData$area |
in \(A_{sj, t}\) | trap area |
mastSim
assumes that predictors of maturation and
fecundity are limited to intercept and diameter. Thus, the
formulaFec
and formulaRep
are identical. Here
is fecundity:
Before fitting the model I say a bit more about input data.
treeData
and
xytree
The information on individual trees is held in the tree-years by
variables data.frame treeData
. Here are a few lines of
treeData
generated by mastSim
,
treeData
has a row for each tree-year. Several of these
columns are required:
data.frame treeData
columns
treeData column |
explanation |
---|---|
plot |
plot name |
tree |
identifier for each tree, unique within a plot |
year |
observation year |
species |
species name |
diam |
diameter is a predictor for fecundity in \(\mathbf{X}\) and maturation in \(\mathbf{V}\) |
repr |
immature ( 0 ), mature ( 1 ),
or unknown ( NA ) |
There can be additional columns, but they are not required for model
fitting. The variable repr
is reproduction status, often
NA
.
There is a corresponding data.frame xytree
that holds
tree locations.
Table 5. data.frame xytree
columns
xytree column |
explanation |
---|---|
plot |
plot name |
tree |
identifier for each tree, unique within a plot |
x, y |
locations in the sample plot |
xytree
has fewer rows than treeData
,
because it is not repeated each year–it assumes that tree locations are
fixed. They are cross-referenced by plot
and
tree
:
##seedData
and
xytrap
Seed counts are held in the data.frame seedData
as
trap-years by seed types. Here are a few lines of
seedData
,
Required columns are in Table 6.
Table 6. data.frame seedData
columns
seedData column |
explanation |
---|---|
plot |
plot name |
trap |
identifier for each trap, unique within a plot |
year |
seed-crop year |
active |
fraction of collecting period that the trap was active |
area |
collection area of trap ( m\(^2\) ) |
acerRubr , … |
seedNames for columns with seed
counts |
Seed trap location data are held in the
data.frame xytrap
.
Table 7. data.frame xytrap
columns
xytrap column |
explanation |
---|---|
plot |
plot name |
trap |
identifier for each trap, unique within a plot |
x, y |
map location |
There is no year in xytrap
, because locations are
assumed to be fixed. If a seed trap location is moved during a study,
simply assign it a new trap name. Then seed counts for years when it is
not present are assigned NA
( missing data ) will be
imputed.
Because they are stochastic, not all simulations from
mastSim
generate data with sufficient pattern to allow
model fitting. I can look at the relationship between tree diameters and
seeds on a map for plot mapPlot
and year
mapYear
:
dataTab <- table( treeData$plot, treeData$year )
w <- which( dataTab > 0, arr.ind=T ) # a plot-year with observations
w <- w[sample( nrow( w ), 1 ), ]
mapYears <- as.numeric( colnames( dataTab )[w[2]] )
mapPlot <- rownames( dataTab )[w[1]]
inputs$mapPlot <- mapPlot
inputs$mapYears <- mapYears
inputs$treeSymbol <- treeData$diam
inputs$SCALEBAR <- T
mastMap( inputs )
Depending on the numbers and sizes of maps I adust
treeScale
and trapScale
to see how seed
accumulation compares with tree size or fecundity. In the above map,
trees are shown as green circles and traps as gray squares. The size of
the circle comes from the input variable treeSymbol
, which
is set to tree diameter. I could instead set it to the ‘true’ number of
seeds produced by each tree, an output from mastSim
.
Which are reproductive (mature)? Here are true values, showing just trees that are mature:
To fit the data, there must be sufficient seed traps, and sources must not be so dense such that overlapping seed shadows make the individual contributions undetectable.
Here is the frequency distribution of seeds ( observed ) and of fecundities ( unknown ) from the simulated data:
par( mfrow=c( 1, 2 ), bty='n', mar=c( 4, 4, 1, 1 ) )
seedData <- inputs$seedData
seedNames <- inputs$seedNames
hist( as.matrix( seedData[, seedNames] ) , nclass=100,
xlab = 'seed count', ylab='per trap', main='' )
hist( trueValues$fec, nclass=100, xlab = 'seeds produced', ylab = 'per tree', main = '' )
In data of this type, most seed counts and most tree fecundities are zero.
Model fitting requires specification of the number of MCMC iterations
ng
and the burnin
. Here is an analysis using
the simulated inputs
from mastSim
with a small
number of iterations:
The fitted object output
contains MCMC chains,
estimates, and predictions, summarized in the next section.
output
summary, lists, and
plotsSample size, parameter estimates, goodness of fit are all provides as
tables by summary
.
By default, this summary is sent to the console. It can also be saved
to a list
, e.g.,
outputSummary <- summary( output )
.
output
The main objects returned by mastif
include several
lists summarized in Table 9. parameters
are estimated as
part of model fitting. predictions
are generated by the
fitted model, as predictive distributions. Note that the latent states
for log fecundity \(\psi_{ij, t}\) and
maturation state \(\rho_{ij, t}\) are
both estimated and predicted.
Table 9. The list
created by function
mast
.
list in output |
summary | contents |
---|---|---|
inputs |
from inputs with additions |
includes distall ( trap by tree distance
) |
chains |
MCMC chains | agibbs ( if random effects, the covariance
matrix \(\mathbf{A}\) ),
bfec ( \(\boldsymbol{\beta}^x\) ), brep
( \(\boldsymbol{\beta}^v\) ),
bygibbs ( \(\alpha_l\) or
\(\gamma_t\) if yearEffect
included ), rgibbs ( \(\mathbf{M}\) if multiple seed types ),
sgibbs ( \(\sigma^2\),
RMSPE, deviance ), ugibbs ( \(u\) parameter ) |
data |
data attributes | intermediate objects used in fitting, prediction |
fit |
diagnostics | DIC, RMSPE, scoreStates, predictScore |
parameters |
posterior summaries | alphaMu and alphaSe ( mean
and se for \(\mathbf{A}\), if included
), aMu and aSe ( \(\mathbf{\beta^w}_{ij}\) ),
betaFec ( \(\boldsymbol{\beta}^x\) ),
betaRep ( \(\boldsymbol{\beta}^v\) ),
betaYrMu and betaYrSe ( mean and se for year
or lag coefficients ), upars and dpars (
dispersal parameters \(u\) and \(d\) ), rMu and
rSe ( mean and se for \(\mathbf{M}\) ), sigma ( \(\sigma^2\) ), acfMat ( group
by lag empirical correlation or ACF ), pacfMat ( group by
lag partial correlation or PACF ), pacfSe ( se for
pacfMat ), pacsMat ( group by lag PACF for
seed data ) |
prediction |
predictive distributions | fecPred ( maturation \(\rho_{ij, t}\) and fecundity \(\psi_{ij, t}\) estimates and predictions ),
seedPred ( seed counts per trap and predictions per m\(^2\) ), seedPredGrid
predictions for seeds on the space-time prediction grid,
treePredGrid predictions for maturation and fecundity
corresponding to seedPredGrid . |
Prediction scores for seed-trap observations are provided in
output$fit
based on the estimated fecundities \([\mathbf{y} | \phi, \rho]\) as
scoreStates
and on the estimated parameters \([\mathbf{y} | \phi, \rho][\phi, \rho|
\boldsymbol{\beta}^x, \boldsymbol{\beta}^w, \dots]\) as
predictScore
. The former will be substantially higher than
the latter in cases where estimates of states \(\phi, \rho\) can be found that predict seed
data, but the variables in \(\mathbf{X},
\mathbf{V}\) do not predict those states well. These proper
scoring rules are bases on the log likelihood for the Poisson
distribution.
Predictions for a location-year prediction grid are invoked when
predList
is specified in the call to
mastif
.
output
Here are plots of output
, with the
list plotPars
passing the trueValues
for these
simulated data:
Because this is a simulated data set, I pass the
trueValues
in the list plotPars
.
By inspecting chains I decide that it is not converged and continue,
with output
from the previous fit being the new
inputs
:
Other arguments passed to mastPlot
in the
list plotPars
are given as examples below and listed at
help( mastPlot )
.
mastPlot
generates the following plots:
maturation
: chains for maturation parameters in
\(\beta^v\).
fecundity
: chains for fecundity parameters in \(\beta^x\).
dispersal parameter
: chains for dispersal parameter
\(u\) showing prior
distribution.
variance sigma
: chains for error variance \(\sigma^2\), RMSPE, and deviance.
maturation, fecundity
: posterior 68% ( boxes ) and
95% ( whiskers ) for \(\beta^x\) and
\(\beta^v\).
maturation, fecundity by diameter
: estimates ( dots
) with \(95%\) predictive means for
latent states.
seed shadow
: with 90% predictive interval
prediction
: seed data predicted from estimates of
latent maturation and fecundity ( a ) and from parameter estimates. If
trueValues
are supplied from mastSim
, then
panel ( c ) includes true versus predicted values. There is an important
distinction between ( a ) and ( b )–good predictions in ( a ) indicate
that combinations of seed sources can be found to predict the seed data,
without necessarily meaning that the process model can predict
maturation and fecundity. Good predictions in ( b ) face the steeper
challenge that the process model must predict both maturation and
fecundity, which, in turn, predict seed rain.
parameter recovery
: if trueValues
are
supplied from mastSim
, then this plot is provided comparing
true and estimated values for \(\beta^v\) and \(\beta^x\).
predicted fecundity, seed data
: maps show predicted
fecundity of trees ( sizes of green circles ) with seed data ( gray
squares ). If predList
is supplied to mastif
,
then the predicted seed surface is shown as shaded contours.
partial ACF
: autocorrelation function for fecundity
( estimated ) ( a ) and in seed counts ( observed ) ( b ).
tree correlation in time
: pairwise correlations
between trees on the same plot.
The plots displayed by mastPlot
include the MCMC chains
that are not yet converged. Again, I can restart where I left off by
using output
as the inputs
to
mast
. In addition, I predict the seed surface from the
fitted model for a plot and year, as specified in
predList
.
output$predList <- list( mapMeters = 3, plots = mapPlot, years = mapYears )
output <- mastif( inputs = output, ng = 2000, burnin = 1000 )
To look closer at the predicted plot-year I generate a new map:
output$mapPlot <- mapPlot
output$mapYears <- mapYears
output$treeScale <- 1.2
output$trapScale <- .7
output$PREDICT <- T
output$scaleValue <- 10
output$plotScale <- 1
output$COLORSCALE <- T
output$LEGEND <- T
mastMap( output )
The maps show seed counts ( squares ) and fecundity predictions ( circles ). For predicted plot years there is also shown a seed prediction surface. The surface is seeds per m\(^2\).
Depending on the simulation, convergence may require more iterations.
The partial autocorrelation for years should be weak, because none are
simulated in mastSim
. However, actual data will contain
autocorrelation. The fecundity-time correlations show modal values near
zero, because simulated data do not include individual covariance.
Positive spatial covariance is imposed by dispersal.
To send output to a single Rmarkdown
file and
html
or pdf
, I use this:
This option will generate a file mastifOutput.Rmd
, which
can be opened in Rstudio, edited, and knitted to pdf
format. It contains data and posterior summaries generated by
summary
and mastPlot
. This pdf will not
include stand maps. Maps will be included in the html
version, obtained by setting plotPars$RMD <- 'html'
.
Seed shadow models confront convoluted likelihood surfaces, in the sense that we expect local optima. These surfaces are hard to traverse with MCMC ( and impossible with HMC ), because maturation status is a binary state that must be proposed and accepted together with latent fecundity. Posterior simulation can get bogged down when fecundity estimates converge for a combination of mature and immature trees that is locally but not globally optimal. Especially when there are more trees of a species than there are seed traps, we expect many iterations before the algorithm can ‘find’ the specific combination of trees that together best describe the specific combination of seed counts in many seed traps. Compounding the challenge is the fact that, because both maturation and fecundity are latent variables for each individual, the redistribution kernel must be constructed anew at each MCMC step. Yet, analysis of simulated data shows that convergence to the correct posterior distribution is common. It just may depend on finding reasonable prior parameter values ( see below ) and long chains.
Examples in this vignette assign enough interations to show this
progress toward convergence may be happening, but sufficiently few to
avoid long waiting times. To evaluate convergence, consider the plots
for chains of sigma
( the residual variance \(\sigma\) on log fecundity ), the
rspse
( the seed count residual ), and the
deviance
. Finally, the plot of seed observed vs predicted
gives a sense of progress.
As demonstrated above, restart mastif
with the fitted
object.
Often a seed type could have come from trees of more than one
species. Seeds that are only identified to genus level include in
seedNames
the character
string
UNKN
. In this simulation the three species
pinuTaeda
, pinuEchi
, and pinuVirg
contribute most seeds to the type pinuUNKN
.
Important: there can be only one element in
seedNames
containing the string UNKN
.
Here are some inputs for the simulation:
specNames <- c( 'pinuTaeda', 'pinuEchi', 'pinuVirg' )
seedNames <- c( 'pinuTaeda', 'pinuEchi', 'pinuVirg', 'pinuUNKN' )
sim <- list( nyr=4, ntree=25, nplot=10, ntrap=50, specNames = specNames,
seedNames = seedNames )
There is a specNames
-by-seedNames
matrix
M
that is estimated as part of the posterior distribution.
For this example seedNames
containing the string
UNKN
refers to a seed type that cannot be differentiated
beyond the level of the genus pinu
.
Here is the simulation with output objects with 2/3 of all seeds identified only to genus level:
inputs <- mastSim( sim ) # simulate dispersal data
R <- inputs$trueValues$R # species to seedNames probability matrix
round( R, 2 )
The matrix M
stacks the length-\(M\) vectors \(\mathbf{m}'_s\) discussed in Clark et
al. ( 2019 ) as a species-by-seed type matrix. There is a matrix for
each plot, here stacked as a single matrix. This is the matrix of values
used in simulation. mastif
will estimate this matrix as
part of the posterior distribution.
Here is a model fit:
The model summary now includes estimates for the unknown
M
as the “species to seed type matrix”:
Output plots will include chains for estimates in M
.
There will also be estimates for each species included in
specNames
:
The inverse of \(\mathbf{M}\) is the
probability that an unknown seed of type \(m\) was produced by a tree of species \(s\). These estimates are included in the
plot Species to undiff seed
.
Again, the .Rmd
file can be knitted to a
pdf
file.
Here is a restart, now with plots
and years
specified for prediction in predList
:
tab <- with( inputs$seedData, table( plot, year ) )
mapPlot <- 'p1'
mapYears <- as.numeric( colnames( tab )[tab[1, ] > 0] ) # years for 1st plot
output$predList <- list( plots = mapPlot, years = mapYears )
output <- mastif( inputs = output, ng = 3000, burnin = 1500 )
mastPlot( output, plotPars = list( trueValues = inputs$trueValues, MAPS = TRUE ) )
Chains for the matrix M
will converge for plots where
there are sufficient trees and seeds to obtain an estimate. They will
not be identified on plots where one or the other are rare or
absent.
To map just the seed rain prediction maps, do this:
output$PREDICT <- T
output$LEGEND <- T
output$mapPlot <- mapPlot
output$mapYears <- mapYears
mastMap( output )
specNames
and
seedNames
In the many data sets where seeds of multiple species are not
differentiated the undifferentiated seed type occupies a column in
seedData
having a column name that includes the string
UNKN
. For a concrete example, if specNames
on
a plot include trees in treeData$species
with the
specNames pinuTaed
and pinuEchi
, then many
seeds might be indentified as seedNames pinuUNKN
. In the
foregoing example, estimates of the matrix M
reflect the
contributions of each species to the UNKN
type. This could
be specified in several ways:
mastPlot
written to
filesThe function mastPlot
generates summary plots of
output
. The user has access to all objects used to generate
these plots, as discussed in the previous section. By default plots in
mastPlot
can be rendered to the console. If
plotPars$SAVEPLOTS = T
is included in the
list plotPars
passed to mastPlot
, then each
plot is saved to a .pdf
file. Plots can be consolidated
into a single .Rmd
file, which can be knitted to
.html
or .pdf
with
plotPars$RMD = html
or plotPars$RMD = pdf
.
For illustration I use sample data analyzed by Clark et al. ( 2013 ), with data collection that has continued through 2017. It consists of multiple years and sites.
Here I load data for species with a single recognized seed type, Liriodendron tulipifera. Here is a map, with seed traps scaled by seed counts, and trees scaled by diameter,
library( repmis )
d <- "https://github.com/jimclarkatduke/mast/blob/master/liriodendronExample.rData?raw=True"
repmis::source_data( d )
mapList <- list( treeData = treeData, seedData = seedData,
specNames = specNames, seedNames = seedNames,
xytree = xytree, xytrap = xytrap, mapPlot = 'DUKE_BW',
mapYears = 2011:2014, treeSymbol = treeData$diam,
treeScale = .7, trapScale = 1.5, plotScale = 2,
SCALEBAR=T, scaleValue=50 )
mastMap( mapList )
Here are a few lines of treeData
, which has been trimmed
down to this single species,
Here are a few lines of seedData
,
Here I fit the model.
formulaFec <- as.formula( ~ diam ) # fecundity model
formulaRep <- as.formula( ~ diam ) # maturation model
inputs <- list( specNames = specNames, seedNames = seedNames,
treeData = treeData, seedData = seedData, xytree = xytree,
xytrap = xytrap )
output <- mastif( inputs, formulaFec, formulaRep, ng = 3000, burnin = 1000 )
Following this short sequence, I fit a longer one and predict one of the plots:
output$predList <- list( mapMeters = 10, plots = 'DUKE_EW', years = 2010:2015 )
output <- mastif( inputs = output, ng = 4000, burnin = 1000 )
Here are some plots followed by comments on display panels. Notice that when it progresses to the maps for plot DUKE_HW, they will include the predicted seed shadows:
From fecundity
chains, still more iterations are needed
for convergence.
From seed shadow
, seed rain beneath a 30-cm diameter
tree averages near 0.2 seeds per m\(^2\).
Although a combination of maturation/fecundity can be found to
predict the seed data in prediction
( part a ), the process
model does not well predict the maturation/fecundity combination ( part
b ). In other words, the maturation/fecundity/dispersal aspect of the
model is effective, whereas the design does not yet include variables
that help explain maturation/fecundity.
From the many maps in predicted fecundity, seed data
,
note that the DUKE_EW
site includes prediction surfaces, as
specfied in predList
.
Here is the summary:
At this point in the the Gibbs sampler, the DIC and root mean square prediction error are:
As mentioned above, prediction scores for seed-trap observations are
based on the estimated fecundities \([\mathbf{y} | \phi, \rho]\) as
scoreStates
and on the estimated parameters \([\mathbf{y} | \phi, \rho][\phi, \rho|
\boldsymbol{\beta}^x, \boldsymbol{\beta}^w, \dots]\) as
predictScore
. The TGc
and TGd
parameters are the affine-transformation parameters ( intercept, slope )
relating the predictive variance to the error variance ( Thorarinsdottir
and Gneiting, 2010 ).
Individual ( tree differences ) can be random and year-to-year
differences can be fixed; the latter are not ‘exchangeble’. In cases
where there are no predictors that explain variation among individuals
the formula
can be limited to an intercept, using
~ 1
. In this case, it can be valuable to estimate fecundity
even when there are no good predictors. For this intercept-only model,
random effects and/or year effects can allow for variation. Prior
parameter values are supplied as discussed below.
#group plots in regions for year effects
region <- rep( 'sApps', nrow( treeData ) )
region[ as.character( treeData$plot ) == 'DUKE_EW' ] <- 'piedmont'
treeData$region <- region
formulaFec <- as.formula( ~ diam )
formulaRep <- as.formula( ~ diam )
yearEffect <- list( groups = 'region' )
randomEffect <- list( randGroups = 'treeID', formulaRan = as.formula( ~ 1 ) )
inputs <- list( specNames = specNames, seedNames = seedNames,
treeData = treeData, seedData = seedData,
xytree = xytree, xytrap = xytrap,
priorDist = 28, priorVDist = 15, maxDist = 50, minDist = 15,
minDiam = 25, maxF = 1e+6,
randomEffect = randomEffect, yearEffect = yearEffect )
output <- mastif( inputs, formulaFec, formulaRep, ng = 2000, burnin = 1000 )
mastPlot( output )
Without predictors, the fitted variation is coming from random effects and year effects.
To substitute year effects for AR( \(p\) ) effects, simply remove the argument
specification of p
in the yearEffect list
:
In previous examples, the prior parameter values were past in the
list inputs
. I can also specify prior values as inputs
through the inputs$priorList
or
inputs$priorTable
holding parameters named in Table 8.
Table 8. Components of inputs
with
default values.
object | explanation |
---|---|
priorDist = 25 |
prior mean dispersal distance parameter ( \(m\) ) |
priorVDist = 40 |
prior variance dispersal parameter |
minDist = 4 |
minimum mean dispersal distance ( \(m\) ) |
maxDist = 70 |
maximum mean dispersal distance ( \(m\) ) |
maxF = 1e+8 |
maximum fecundity ( seeds per tree-year ) |
minDiam = 10 |
minimum diam below which a tree can mature
( \(cm\) ) |
maxDiam = 40 |
maximum diam above which a tree can be
immature ( \(cm\) ) |
sigmaMu = 1 |
prior mean residual variance \(\sigma^2\) |
sigmaWt = sqrt( nrow( treeData ) ) |
weight on prior mean ( no. of observations ) |
The mean and variance of the dispersal kernel, priorDist
and priorVDist
, refer to the dispersal parameter \(u\), transformed to the mean distance for
the dispersal kernel. The parameters minDist
and
maxDist
place minimum and maximum bounds on \(d\).
Fecundity \(\phi\) has a prior maximum
value of maxF
.
minDiam
and maxDiam
bound diameter ranges
where a tree of unknown maturation status can be mature or not. This
prior range is overridden by values in the treeData$repr
column that establish an observed maturation state for a tree-year ( 0 -
immature, 1 - mature ).
In general, large-seeded species, especially those that can be
dispersed by vertebrates, generate noisy seed-trap data, despite the
fact that the bulk of the counts still occur close to the parent, and
the mean dispersal distance is relatively low. Large-seeded species
produce few fruits/seeds. Long-distance dispersal cannot be estimated
from inventory plots, regardless of plot size, because the fit is
dominated by locally-derived seed. Consider values for maximum fecundity
as low as maxF = 10000
, minimum dispersal
minDist = 2
, and maximum dispersal
maxDist = 12
. Recall that the latter values refer to the
dispersal kernel parameter value, not the maximum distance a seed can
travel, which is un-constrained.
Prior parameter values can be passed as a list
,
e.g.,
Alternatively, prior parameter values can be passed as a table, with
parameters for column names for specNames
for rownames.
Here is a file holding parameter values and the
function mastPriors
to obtain a table for several species
in the genus \(Pinus\),
d <- "https://github.com/jimclarkatduke/mast/blob/master/priorParametersPinus.csv?raw=True"
download.file( d, destfile="priorParametersPinus.csv" )
specNames <- c( "pinuEchi", "pinuRigi", "pinuStro", "pinuTaed", "pinuVirg" )
seedNames <- c( "pinuEchi", "pinuRigi", "pinuStro", "pinuTaed", "pinuVirg", "pinuUNKN" )
priorTable <- mastPriors( "priorParametersPinus.csv", specNames,
code = 'code4', genus = 'pinus' )
The argument code = 'code8'
specifies the column in
priorParametersPinus.csv
holding the codes corresponding to
specNames
. In mastif
the format
code4
means that names consist of the first 4 letters from
the genus and species, like this: pinuTaed
. The
genus
is provided to allow for genus-level parameters in
cases where priors for individual species have not been specified in
priorParametersPinus.csv
.
For coefficients in fecundity, betaPrior
specifies
predictor effects by sign. The use of prior distributions that are flat,
but truncated, is two-fold ( Clark et al. 2013 ). First, where prior
information exists, it often is limited to a range of values. For
regressions coefficients ( e.g., \(\boldsymbol{\beta}\) ) we typically have a
prior belief about the sign of the effect ( positive or negative ), but
not its magnitude or appropriate weight. Second, within bounds placed by
the prior, the posterior has the shape of the likelihood, unaffected by
a prior weight. Prior weight is hard to specify sensibly for this
hierarchical, non-linear model.
Here is an example using betaPrior
to specify a
quadratic diameter effect. First, here is a data set for
Pinus:
d <- "https://github.com/jimclarkatduke/mast/blob/master/pinusExample.rdata?raw=True"
repmis::source_data( d )
Here I specify formulas, prior bounds for diam
(
quadratic ), and fit the model:
formulaRep <- as.formula( ~ diam )
formulaFec <- as.formula( ~ diam + I( diam^2 ) )
betaPrior <- list( pos = 'diam', neg = 'I( diam^2 )' )
inputs <- list( treeData = treeData, seedData = seedData, xytree = xytree,
xytrap = xytrap, specNames = specNames, seedNames = seedNames,
betaPrior = betaPrior, priorTable = priorTable,
seedTraits = seedTraits )
output <- mastif( inputs = inputs, formulaFec, formulaRep,
ng = 500, burnin = 200 )
# restart
output <- mastif( output, formulaFec, formulaRep, ng = 5000, burnin = 1000 )
mastPlot( output )
Here again, many of the seed ID maxtrix elements are well identified
( trace plots labeled “M matrix plot
”, others are uncertain
due to small numbers of the different species that could contribute to a
seed type. These are presented as bar and whisker plots for elements of
matrix \(\mathbf{M}\) ( fraction of
seed from a species that is counted as each seed type ) in the plot
labeled species -> seed type
. Bar plots show the
fraction of unknown seeds that derive from each species, labeled
species -> seed type
. The distribution of diameters in
the data will limit the ability to estimate its effect on maturation and
fecundity.
Note that the estimates for fecundity coefficients, \(\boldsymbol{\beta}^x\), are positive for
diam
and negative for diam^2
. If there is no
evidence in the data for a decrease \(\frac{\partial{\psi}}{\partial{diam}}\),
then the quadratic term is near zero. In this case, the predicted
fecundity in the plot labelled
maturation, fecundity by diameter
will increase
exponentially.
Most individuals produce no seed, due to resource limitation,
especially light, in crowded stands. As trees increase in size and
resource access they mature, and become capable of seed production.
Maturation is hard to observe, so it must be modelled. In our data sets,
maturation status is observed, but most values are NA; it is only
assigned if certain. A value of 1 means that reproduction is observed. A
value of 0 means that the entire crown is visible during the fruiting
season, and it is clear that no reproduction is present. Needless to
say, most observations from beneath closed canopies are NA. The
observations are entered in the column repr
in the
data.frame treeData
. Above I showed an example where
minDiam
is set as a prior that trees of smaller diameters
are immature and maxDiam
is the prior that trees above this
diameter are mature.
mastif
further admits estimates of cone production. The
column treeData$cropCount
refers to the cones that will
open and contribute to trap counts in the year
treeData$year
. There is another column
treeData$cropFraction
, which refers to the fraction of the
tree canopy that is represented by the cone count. When crop counts are
used, a matrix seedTraits
must be supplied with one row per
species and a column with seeds per fruit, cone, or pod. The rownames of
seedTraits
are the specNames
. The colname of
seedTraits
with seeds per fruit is labeled
seedsPerFruit
.
Seed studies often include seed collections in years when trees are
not censused. For example, tree data might be collected every 2 to 5
years, whereas seed data are available as annual counts. In this case,
there there are years in seedData$year
that are missing
from treeData$year
. mastif
needs a tree year
for each trap year. If tree data are missing from some trap years, I
need to constuct a complete treeData data.frame
that
includes these missing years. This amended version of
treeData
must be accessible to the user, to allow for the
addition of covariates such as weather variables, soil type, and so
forth.
The function mastFillCensus
allows the user to access
the filled-in version of treeData
that will be fitted by
mastif
. mastFillCensus
accepts the same
list
of inputs
that is passed to the function
mastif
. The missing years are inserted for each tree with
interpolated diameters. The list inputs
is returned with
objects updated to include the missing census years and modified
slightly for analysis by mastif
.
inputs$treeData
can now be annotated with the covariates
that will be included in the model. Here is the example from the
help
file. First I read a file that has complete years, so
I randomly remove years:
# randomly remove years for this example:
years <- sort( unique( treeData$year ) )
sy <- sample( years, 5 )
treeData <- treeData[treeData$year %in% sy, ]
treeData[1:10, ]
Note the missing years, as is typical of mast data sets. Here is the file after filling missing years:
inputs <- list( specNames = specNames, seedNames = seedNames,
treeData = treeData, seedData = seedData,
xytree = xytree, xytrap = xytrap, priorTable = priorTable,
seedTraits = seedTraits )
inputs <- mastFillCensus( inputs, beforeFirst=10, afterLast=10 )
inputs$treeData[1:10, ]
The missing plotYr
combinations in treeData
have been filled to match thos in seedData
. Now columns can
be added to inputs$treeData
, as needed in
formulaFec
or formulaRep
.
In cases where tree censuses start after seed trapping begins or tree
censuses end before seed trapping ends, it may be reasonable to assume
that the same trees are producing seed in those years pre-trapping or
post-trapping years. In these cases, mastFillCensus
can
accommodate these early and/or late seed trap data by extrapolating
treeData
beforeFirst
years before seed
trapping begins or afterLast
years after seed tapping
ends.
Continuing with the \(Pinus\) example, here I want to add covariates for missing years as needed for an AR( \(p\) ) model, in this example \(p\) = 4. First, consider the tree-years included in this sample, before and after in-filling:
# original data
table( treeData$year )
# filled census
table( inputs$treeData$year )
table( seedData$year )
Note that the infilled version of tree data in the
list inputs
has the missing years, corresponding to those
included in seedData
. Here I set up the year effects model
and infill to allow for p = 3
,
Here are regions for random effects in the AR( 3 ) model:
region <- c( 'CWT', 'DUKE', 'HARV' )
treeData$region <- 'SCBI'
for( j in 1:length( region ) ){
wj <- which( startsWith( treeData$plot, region[j] ) )
treeData$region[wj] <- region[j]
}
inputs$treeData <- treeData
yearEffect <- list( groups = c( 'species', 'region' ), p = p )
I add the climatic deficit ( monthly precipitation minus PET ) as a covariant. First, here is a data file,
d <- "https://github.com/jimclarkatduke/mast/blob/master/def.csv?raw=True"
download.file( d, destfile="def.csv" )
The format for the file dev.csv
is plot
by
year_month
. I have saved it in my local directory. The
function mastClimate
returns a list
holding
three vectors, each having length equal to
nrow( treeData )
. Here is an example for the cumulative
moisture deficit for the previous summer. I provide the file name, the
vector of plot names, the vector of previous years, and the months of
the year. To get the cumulative deficit, I use
FUN = 'sum'
:
treeData <- inputs$treeData
deficit <- mastClimate( file = 'def.csv', plots = treeData$plot,
years = treeData$year - 1, months = 6:8,
FUN = 'sum', vname='def' )
treeData <- cbind( treeData, deficit$x )
summary( deficit )
The first column in deficit
is the variable itself for
each tree-year. The second column holds the site mean value for the
variable. The third column is the difference between the first two
columns All three can be useful covariates, each capturing different
effects ( Clark et al. 2014 ). I could append any or all of them as
columns to treeData
.
Here is an example using minimum temperature of the preceeding
winter. I obtain the minimum with two calls to mastClimate
,
first for Dec of the previous year (
years = treeData$year - 1, months = 12
), then from Jan,
Feb, Mar for the current year (
years = treeData$year, months = 1:3
). I then take the
minimum of the two values:
# include min winter temperature
d <- "https://github.com/jimclarkatduke/mast/blob/master/tmin.csv?raw=True"
download.file( d, destfile="tmin.csv" )
# minimum winter temperature December through March of previous winter
t1 <- mastClimate( file = 'tmin.csv', plots = treeData$plot,
years = treeData$year - 1, months = 12, FUN = 'min',
vname = 'tmin' )
t2 <- mastClimate( file = 'tmin.csv', plots = treeData$plot,
years = treeData$year, months = 1:3, FUN = 'min',
vname = 'tmin' )
tmin <- apply( cbind( t1$x[, 1], t2$x[, 1] ), 1, min )
treeData$tminDecJanFebMar <- tmin
inputs$treeData <- treeData
Here is a model using several of these variables:
formulaRep <- as.formula( ~ diam )
formulaFec <- as.formula( ~ diam + defJunJulAugAnom + tminDecJanFebMar )
inputs$yearEffect <- yearEffect
output <- mastif( inputs = inputs, formulaFec, formulaRep, ng = 2500, burnin = 1000 )
Data for future years can be a scenario, based on assumptions of status quo in the mean and variance, an assumed rate of climate change, and so on.
Seed production is volatile, with order of magnitude variation from year-to-year. There is synchronicity among individuals of the same species, the “masting” phenomenon. There are large difference between individuals, that are not explained by environmental variables. In this section I discuss extensions to random effects, year effects, lag effects, and fitting multiple species having seed types that are not always identifiable to species.
Random individual effects currently can include random intercepts for the fecundities of each individual tree that is imputed to be in the mature state for at least 3 years. [There are no random effects on maturation, because they would be hard to identify from seed trap data for this binary response.]
I include in the inputs list
the
list randomEffect
, which includes the column name for the
random group. Typically this would be a unique identifier for a tree
within a plot, e.g., randomEffect$randGroups = 'tree'
.
However, randGroups
could be the plot name. This column is
interpreted as a factor
, each level being a group. Random
effects will not be fitted on individuals that are in the mature state
less than 3 years.
The formulaRan
is the random effects model. Because
individual time series tend to short, it is currently implemented only
for intercepts. Here is a fit with random effects.
formulaFec <- as.formula( ~ diam ) # fecundity model
formulaRep <- as.formula( ~ diam ) # maturation model
inputs$randomEffect <- list( randGroups = 'tree', formulaRan = as.formula( ~ 1 ) )
output <- mastif( inputs = inputs, formulaFec, formulaRep, ng = 2000, burnin = 1000 )
Here is a restart:
There is a new panel for fixed plus random effects
,
showing the individual combinations of intercepts.
prediction
panel ( b ) shows some improvement,
indicating that even with random effects, diameter struggles to predict
maturation/fecundity.As discussed previously, the model admits year effects and lag effects, the latter as an AR( \(p\) ) model. Year effects assign a coefficient to each year \(t = [1, \dots, T_j]\). Lag effects assign a coefficient to each of \(j \in \{1, \dots, p\}\) plags, where the maximum lag \(p\) should be substantially less than the number of years in the study. Year effects can be organized in random groups. Specification of random groups is done in the same way for year effects and for lag effects.
I define random groups for year and lag effects by species, by plots,
or both. When there are multiple species that contribute to the modeled
seed types, I expect the year effects to depend on which species is
actually producing the seed. When there are multiple plots sufficiently
distant from one another, I might allow for the fact that year effects
or lag effects differ by group; yearEffect$groups
allows
that they need not mast in the same years. In the example below, I’ll
use the term region for plots in the same plotGroup.
Here is a breakdown for this data set by region:
Here are year effects structured by random groups of plots, given by
the column region
in treeData
:
This option will fit a year effect for both provinces and years having sufficient individuals estimated to be in the mature state. Here is the model with random year effects,
\[\log \psi_{ij, t} \sim N \left(
\mathbf{x}'_{ij, t} \mathbf{\beta}^x + \gamma_t + \gamma_{g[i], t},
\sigma^2 \right )\] where the year effect \(\gamma_{g[i], t}\) is shared by trees in
all plots defined by yearEffect$province
, \(ij \in g\). Year effects are sampled
directly from conditional posteriors.
inputs$treeData <- treeData
inputs$randomEffect <- randomEffect
inputs$yearEffect <- yearEffect
output <- mastif( inputs = inputs, formulaFec, formulaRep, ng = 2500, burnin = 500 )
Here is a restart, with predictions for one of the plots:
predList <- list( mapMeters = 10, plots = 'DUKE_BW', years = 1998:2014 )
output$predList <- predList
output <- mastif( inputs = output, ng = 3000, burnin = 1000 )
mastPlot( output )
Note that still more iterations are needed for convergence. Here are
some comments on mastPlot
:
In the dispersal parameter u
panel there are now
year effects plotted for the two random groups mtn
and
piedmont
.
The subsequent panel dispersal mean and variance
shows the mean and variance of random effects
There is a dispersal by group
panel showing
posterior estimate for the two random groups, with scales for the
parameter \(u\) on the left ( m\(^2\) ) and mean parameter \(d\) on the right ( m ).
There has been some improvement in the prediction
,
panel ( b ).
The predicted fecundity, seed data
maps for the plot
DUKE_BW
show seed prediction surfaces.
The year effect groups
shows year effects for random
groups in treeData$region
.
partial ACF
shows partial autocorrelation by species
and plot.
Most data sets have multiple seed types that complicate estimation of mast production by each species. This example considers Pinus spp, seeds of which cannot be confidently assigned to species. Here I load the data and generate a sample of maps from several years, including all species and seed types.
d <- "https://github.com/jimclarkatduke/mast/blob/master/pinusExample.rdata?raw=True"
repmis::source_data( d )
mapList <- list( treeData = treeData, seedData = seedData,
specNames = specNames, seedNames = seedNames,
xytree = xytree, xytrap = xytrap, mapPlot = 'DUKE_EW',
mapYears = c( 2007:2010 ), treeSymbol = treeData$diam,
treeScale = .6, trapScale=1.4,
plotScale = 1.2, LEGEND=T )
mastMap( mapList )
Note the tendency for high seed accumulation ( large green squares ) near dense, large trees ( large brown circles ).
In this example, I again model seed production as a function of log
diameter, diam
, now for multiple species and seed types.
This is an AR( p ) model, because I include the number of lag terms
yearEffect$p = 5
,
\[\log \psi_{ij, t} \sim N \left( \mathbf{x}'_{ij, t} \mathbf{\beta}^x + \sum^p_{l=1} ( \alpha_l + \alpha_{g[i], l} ) \psi_{ij, t-l}, \sigma^2 \right )\] Only years \(p < t \le T_i\) are used for fitting. Samples are drawn directly from the conditional posterior distribution.
In the table printed at the outset are trees by plot and year, i.e.,
the groups
assigned in inputs$yearEffect
. The
zeros indicate either absence of trees or that no plots were sampled in
those years. ( These are not the same thing, mastif
knows
the difference ).
In the code below I specify formulas, AR( \(p\) ), and random effects, and some prior values. Due to the large number of trees, convergence is slow. Because I do not assume that trees of different species necessarily mast in the same years, I allow them to differ through random groups on the AR( \(p\) ) terms.
formulaFec <- as.formula( ~ diam ) # fecundity model
formulaRep <- as.formula( ~ diam ) # maturation model
yearEffect <- list( groups = 'species', p = 4 ) # AR( 4 )
randomEffect <- list( randGroups = 'tree',
formulaRan = as.formula( ~ 1 ) )
inputs <- list( specNames = specNames, seedNames = seedNames,
treeData = treeData, seedData = seedData,
yearEffect = yearEffect,
randomEffect = randomEffect,
xytree = xytree, xytrap = xytrap, priorDist = 20,
priorVDist = 5, minDist = 15, maxDist = 30,
minDiam = 12, maxDiam = 40,
maxF = 1e+6, seedTraits = seedTraits )
output <- mastif( inputs = inputs, formulaFec, formulaRep, ng = 500, burnin = 100 )
Here is a restart:
output <- mastif( inputs = output, ng = 3000, burnin = 1000 )
mastPlot( output, plotPars = plotPars )
Again, convergence will require more iterations. The AR( \(p\) ) coefficients in the
lag effect group
panel shows the coefficients by random
group. They are also shown in a separate panel, with each group plotted
separately. In the ACF eigenvalues
panel are shown the
eigenvalues for AR lag coefficients on the real ( horizontal ) and
imaginary ( vertical ) scales with the unit circle, within which
oscillations are damped. The imaginary axis describes oscillations.
Here’s a restart with predictions:
plots <- c( 'DUKE_EW', 'CWT_118' )
years <- 1980:2025
output$predList <- list( mapMeters = 10, plots = plots, years = years )
output <- mastif( inputs = output, ng = 3000, burnin = 1000 )
and updated plots:
Note that convergence requires additional iterations ( larger
ng
). The predictions of seed production will progressively
improve with convergence.
mapList <- output
mapList$mapPlot <- 'DUKE_EW'
mapList$mapYears <- c( 2011:2012 )
mapList$PREDICT <- T
mapList$treeScale <- .5
mapList$trapScale <- .8
mapList$LEGEND <- T
mapList$scaleValue <- 50
mapList$plotScale <- 2
mapList$COLORSCALE <- T
mapList$mfrow <- c( 2, 1 )
mastMap( mapList )
Or a larger view of a single map:
mapList$mapPlot <- 'CWT_118'
mapList$mapYears <- 2015
mapList$PREDICT <- T
mapList$treeScale <- 1.5
mapList$trapScale <- .8
mapList$LEGEND <- T
mapList$scaleValue <- 50
mapList$plotScale <- 2
mapList$COLORSCALE <- T
mapList$mfrow <- c( 1, 1 )
mastMap( mapList )
Here is a summary of parameter estimates:
R code is highly vectorized. Unavoidable loops are written in C++ and
exploit the C++ library Armadillo
, available through
RcppArmadillo
. Alternating with Metropolis are Hamiltonian
MC steps to encourage large movements.
Despite extensive vectorization and C++ for cases where loops are unavoidable, convergence can be slow. A collection of plots inventoried over dozens of years can generate in excess of \(10^6\) tree-year observations and \(10^4\) trap year observations. There is no escaping the requirement of large numbers of indirectly sampled latent variables. If random effects are included, there are ( obviously ) as many random groups as there are trees. All tree fecundities must be imputed.
Qui et al. (2023) introduced volatility and period to characterize and compare masting behavior between trees and species. The tradition coefficient of variation (CV) ignores the time-dependence in quasi-periodic seed production. To avoid confusion with indices based on variance, they introduced the term volatility as the period-weighted spectral density, to allow for the fact that long intervals are especially important for masting causes and effects on consumers. In addition to this period-weighting of spectral variance within a series (a tree), fecundity-weighting is important at the population scale, because the highly productive individuals dominate masting effects. Within this framework, periodicity extracts the period (in years) that is likewise weighted to emphasize variance concentrated at long intervals within trees and fecundity differences between trees.
The block of code that follows loads output from some trees in the
genus Abies from western North America. Specifically, the
estimates for fecundity by tree-year come from the object
output$prediction$fecPred
. it uses the function
mastVolatility
to compile volatility and periodicity on
each tree and display summaries by ecoregion-species as density plots
and as distributions of mean period estimates.
First, several more functions:
getColor <- function( col, trans ){ # transparent colors
tmp <- col2rgb( col )
rgb( tmp[ 1, ], tmp[ 2, ], tmp[ 3, ], maxColorValue = 255,
alpha = 255*trans, names = paste( col, trans, sep = '_' ) )
}
getPlotLayout <- function( np, WIDE = TRUE ){
# np - no. plots
if( np == 1 )return( list( mfrow = c( 1, 1 ), left = 1, bottom = c( 1, 2 ) ) )
if( np == 2 ){
if( WIDE )return( list( mfrow = c( 1, 2 ), left = 1, bottom = c( 1, 2 ) ) )
return( list( mfrow = c( 2, 1 ), left = c( 1, 2 ), bottom = 2 ) )
}
if( np == 3 ){
if( WIDE )return( list( mfrow = c( 1, 3 ), left = 1, bottom = c( 1:3 ) ) )
return( list( mfrow = c( 3, 1 ), left = 1:3, bottom = 3 ) )
}
if( np <= 4 )return( list( mfrow = c( 2, 2 ), left = c( 1, 3 ), bottom = c( 3:4 ) ) )
if( np <= 6 ){
if( WIDE )return( list( mfrow = c( 2, 3 ), left = c( 1, 4 ), bottom = c( 4:6 ) ) )
return( list( mfrow = c( 3, 2 ), left = c( 1, 3, 5 ), bottom = 5:6 ) )
}
if( np <= 9 )return( list( mfrow = c( 3, 3 ), left = c( 1, 4, 7 ), bottom = c( 7:9 ) ) )
if( np <= 12 ){
if( WIDE )return( list( mfrow = c( 3, 4 ), left = c( 1, 5, 9 ), bottom = c( 9:12 ) ) )
return( list( mfrow = c( 4, 3 ), left = c( 1, 4, 7, 10 ), bottom = 10:12 ) )
}
if( np <= 16 )return( list( mfrow = c( 4, 4 ), left = c( 1, 5, 9, 13 ),
bottom = c( 13:16 ) ) )
if( np <= 20 ){
if( WIDE )return( list( mfrow = c( 4, 5 ), left = c( 1, 6, 11, 15 ),
bottom = c( 15:20 ) ) )
return( list( mfrow = c( 5, 4 ), left = c( 1, 5, 9, 13 ), bottom = 17:20 ) )
}
if( np <= 25 )return( list( mfrow = c( 5, 5 ), left = c( 1, 6, 11, 15, 20 ),
bottom = c( 20:25 ) ) )
if( np <= 25 ){
if( WIDE )return( list( mfrow = c( 5, 6 ), left = c( 1, 6, 11, 15, 20, 25 ),
bottom = c( 25:30 ) ) )
return( list( mfrow = c( 6, 5 ), left = c( 1, 6, 11, 16, 21, 26 ), bottom = 26:30 ) )
}
if( np <= 36 ){
return( list( mfrow = c( 6, 6 ), left = c( 1, 7, 13, 19, 25, 31 ), bottom = c( 31:36 ) ) )
}
return( list( mfrow = c( 7, 6 ), left = c( 1, 7, 13, 19, 25, 31, 37 ), bottom = c( 37:42 ) ) )
}
plotFec <- function( fec, groups = NULL, LOG = F ){
if( is.null(groups) ){
groupID <- rep(1, nrow(fec) )
ngroup <- 1
}else{
group <- fec[, groups]
groups <- sort( unique( group ) )
groupID <- match( group, groups )
ngroup <- length( groups )
}
if( LOG )fec$fecEstMu <- log10(fec$fecEstMu)
xlim <- range( fec$year )
ylim <- range( fec$fecEstMu, na.rm = T )
mfrow <- getPlotLayout(ngroup)
par( mfrow = mfrow$mfrow, bty = 'n', mar = c(4,4,1,1), omi = c( .5, .5, .2, .2) )
for( k in 1:ngroup ){
fk <- fec[ groupID == k, ]
tree <- fk$treeID
trees <- sort( unique( tree ) )
ntree <- length( trees )
plot( NA, xlim = xlim, ylim = ylim, xlab = '', ylab = '' )
for(j in 1:ntree){
fj <- fk[ tree == trees[j], ]
lines( fj$year, fj$fecEstMu, lwd = 1 )
}
title( groups[k] )
}
mtext( 'Year', 1, outer = T, cex = 1.2 )
ytext <- 'Seeds per tree'
if( LOG ) ytext <- 'Seeds per tree (log_10)'
mtext( ytext, 2, outer = T, cex = 1.2 )
}
Here is the volatility:
d <- "https://github.com/jimclarkatduke/mast/blob/master/outputAbies.rdata?raw=True"
repmis::source_data( d )
specs <- sort( unique( fecPred$species ) ) # accumulate period estimates
yseq <- seq( 0, 10, length = 100 )
intVal <- matrix( 0, length(specs), 100 )
weight <- intVal
rownames( intVal ) <- specs
plotSpecs <- sort( unique( fecPred$plotSpec ) ) # label trees in a plot-species group
# time series for one species:
plotFec( fec = fecPred[ fecPred$species == 'abiesGrandis', ], groups = 'species', LOG = T )
par( mfrow = c(1, 2), bty = 'n', mar = c(3,4,1,1), omi = c(.5,.1,.1,.1) )
plot( NA, xlim = c(1, 20), ylim = c(.01, 1), xlab = '', ylab = 'Density/nyr', log = 'xy')
for( i in 1:length(plotSpecs) ){
wi <- which( fecPred$plotSpec == plotSpecs[i] ) # tree-years in group
ci <- fecPred$species[wi[1]]
col <- match( ci, specs ) # color by species
tmp <- mastVolatility( treeID = fecPred$treeID[wi], year = fecPred$year[wi],
fec = fecPred$fecEstMu[wi] )
if( is.null(tmp) )next
intVal[ col, ] <- intVal[ col, ] + dnorm( yseq, tmp$stats['Period', 1], tmp$stats['Period', 2] )
weight[ col, ] <- weight[ col, ] + length( wi )
# density +/- 1 SD
dens <- tmp$statsDensity
lines( dens[ 'Period', ], dens[ 'Mean', ], lwd = 2, col = getColor( col, .4) )
lines( dens[ 'Period', ], dens[ 'Mean', ] - dens[ 'SD', ], lty = 2, col = getColor( col, .4) )
lines( dens[ 'Period', ], dens[ 'Mean', ] + dens[ 'SD', ], lty = 2, col = getColor( col, .4) )
}
title( 'a) Plot-species groups' )
legend( 'topright', specs, text.col = c(1:length(specs)), bty = 'n', cex = .8 )
intVal <- intVal*weight/weight
plot( NA, xlim = c(2, 8), ylim = c(0, .12), xlab = '', ylab = 'Density' ) # density of mean intervals
for( i in 1:length(specs) ){
polygon( yseq, intVal[i,]/sum(intVal[i,]), border = i, col = getColor(i, .4) )
}
title( 'b) Period estimates' )
mtext( 'Year', 1, outer = T, cex = 1.3 )
The following block of code compiles the same densities and summaries
for year effects, which remain after accounting for other predictors in
the model. When fitted with year effects, with
yearEffect <- list( groups = c( 'species', 'ecoCode' ) )
,
the column treeData$ecoCode
holds the ecoregion where the
tree lives. The ecoRegion_species
combinations represent
random groups of year effects that are shared by all trees of the same
species within the same ecoregion. These combinations are
rownames
in output$parameters$betaYrRand
. Here
I use the function mastSpectralDensity
to evaluate
individual rows in betaYrRand
.
spec <- strsplit( rownames( betaYrRand ), '_' ) # extract species from ecoregion_species rownames
spec <- sapply( spec, function(x) x[2] )
specs <- sort( unique(spec) )
mastMatrix <- matrix( 0, nrow(betaYrRand), 5 ) # store stats by group
rownames(mastMatrix) <- rownames(betaYrRand)
colnames(mastMatrix) <- c( 'nyr', 'Variance', 'Volatility', 'Period Est', 'Period SD' )
plot( NA, xlim = c(2, 10), ylim = c(.002, .4), xlab = '', ylab = 'Density/yr', log = 'xy')
for(i in 1:nrow(betaYrRand)){
wc <- which( betaYrRand[i,] != 0 )
if( length(wc) < 6 )next
s <- mastSpectralDensity( betaYrRand[i,wc] )
if( !is.matrix( s$spect ) )next
mastMatrix[i, ] <- c( length(wc), s$totVar, s$volatility, s$periodMu, s$periodSd )
period <- 1/s$spec[, 'frequency' ]
dens <- s$spec[, 'spectralDensity' ]/length(wc) # series vary in length
col <- match( spec[i], specs )
lines( period, dens, lwd = 2, col = getColor( col, .4) )
}
title( 'c) Year effects' )
mtext( 'Period (yr)', 1, line = 1, outer = T )
keepRows <- which( is.finite(mastMatrix[,'Variance']) & mastMatrix[,'Variance'] != 0 )
#keepCols <- which( colSums( frequency, na.rm=T ) > 0 )
mastMatrix <- mastMatrix[ keepRows, ]
Because seed-trap studies involve multiple data sets ( seed traps,
trees, covariates ) that are collected over a number of years and
multiple sites, combining them can expose inconsistencies that are not
immediately evident. Of course, a proper analysis depends on alignment
of trees, seed traps, and covariates with unique tree names (
treeData$tree
) and trap names ( seedData$trap
) in each plot ( treeData$plot, seedData$plot
).
Notes are displayed by mastif
at execution summarizing
aspects of the data that might trigger warnings. All of these issues
have arisen in data sets I have encountered from colleagues:
Alignment of data frames. The unique trees in each
plot supplied in treeData
must also appear with
x
and y
in xytree
. The unique
traps in each plot supplied in seedData
must also appear
with x
and y
in xytrap
. Problems
generate a note:
Note: treeData includes trees not present in xytree
Spatial coordinates. Because tree censuses and seed
traps are often done at different times, by different people, the grids
often disagree. Spatial range
tables are displayed for ( x,
y ) coordinates in xytree
and xytrap
.
Unidentified seeds. Seeds that cannot be identified
to species contain the character
string UNKN
.
If there are species in specNames
that do not appear in
seedNames
, then the UNKN
seed type must be
included in seedNames
and in columns of
seedData
. For example, if caryGlab, caryTome
appear in specNames
, and caryGlab, caryUNKN
appear in seedNames
( and as columns in
seedData
), then caryUNKN
will be the imputed
fate for all seeds emanating from caryOvat
and some seeds
from caryGlab
.
This note will be displayed:
Note: unknown seed type is caryUNKN
If there are seedNames
that do not appear in
specNames
, this note is given:
Note: seedNames not in specNames and not "UNKN":
caryCord
Moved caryCord to "UNKN" class
Design issues. The design can seem confusing, because there are multiple species on multiple plots in multiple years. There is a design matrix that can be found here for fecundity:
output$inputs$setupData$xfec
and here for maturation:
output$inputs$setupData$xrep
( Variables are standardized, because fitting is done that way, but
coefficients are reported on their original scales. Unstandardized
versions of design matrices are xfecU
and
xrepU
. )
There should not be missing values in the columns of
treeData
that will be used as predictors ( covariates or
factors ). If there are missing values, a note will be generated:
Fix missing values in these variables:
[1] "yearlyPETO.tmin1, yearlyPETO.tmin2, flowering.covs.pr.data, flowering.covs.tmin.data, s.PETO"
There can be missing seed counts in seedData
–missing
values will be imputed.
A table containing the Variance Inflation Factor ( VIF ), range of
each variable, and correlation matrix will be generated at execution.
VIF values > 10 and high correlations between covariates are taken as
evidence of redundancy. A table will be generated for each species
separately. However, in xfec
and xrep
they are
treated as a single matrix.
Year effects by random group require replication within groups. Here
is a note for the AR model showing sizes of groups defined by species
and region ( NE, piedmont, sApps
):
no. trees with > plag years, by group:
`caryGlab-NE caryGlab-piedmont caryGlab-sApps `
` 1 497 361 `
`caryOvat-piedmont caryTome-piedmont caryTome-sApps `
` 122 569 77 `
small group: caryGlab-NE
There is only one tree in the caryGlab-NE
group,
suggesting insufficient replication and a different aggregation
scheme.
For the AR( \(p\) ) model, values
are imputed for \(p\) years before and
after a tree is observed, and only trees observed for > \(p\) years will contribute to parameter
estimates. If the study lasts 3 years, then the model should not specify
yearEffect$p = 5
. A note will be generated to inform on the
number of observations included in parameter estimates:
Number of full observations with AR model is: [1] 21235
Prediction. If predList
is supplied,
then fecundity and seed density will be predicted for specified
plot-years. The size of the prediction grid is displayed as a table of
prediction nodes by plot and year. Large prediction grids slow
execution. To reduce the size of the grid, increase the
inputs$predList$mapMeters
( the default is 5 m by 5 m
).
When covariates are added as columns to treeData
, they
must align with treeData$plot
, treeData$year
,
and, if they are tree-level covariates, with treeData$tree
.
The required column treeData$diam
is an example of the
latter.
Qiu T., et al., 2022. Limits to reproduction and seed size-number tradeoffs that shape forest dominance and future recovery. Nature Communications, 13:2381 | https://doi.org/10.1038/s41467-022-30037-9
Journe, V., et al., 2022. Globally, tree fecundity exceeds productivity gradients. Ecology Letters, DOI: 10.1111/ele.14012, pdf: Ecology Letters2022.
Sharma, S., et al., 2022. North American tree migration paced by recruitment through contrasting east-west mechanisms. Proceedings of the National Academy of Sciences, 119 (3) e2116691118; https://doi.org/10.1073/pnas.2116691118.
Qiu, T., et al., 2021. Is there tree senescence? The fecundity evidence. Proceedings of the National Academy of Sciences, 118, e2106130118; DOI: 10.1073/pnas.2106130118.
Clark, J.S., 2021. Continent-wide tree fecundity driven by indirect climate effects. Nature Communications DOI: 10.1038/s41467-020-20836-3.
Clark, JS, C Nunes, and B Tomasek. 2019. Masting as an unreliable resource: spatio-temporal host diversity merged with consumer movement, storage, and diet. Ecological Monographs, 89:e01381.
Clark, JS, S LaDeau, and I Ibanez. 2004. Fecundity of trees and the colonization-competition hypothesis, Ecological Monographs, 74, 415-442.
Clark, JS, M Silman, R Kern, E Macklin, and J Hille Ris Lambers. 1999. Seed dispersal near and far: generalized patterns across temperate and tropical forests. Ecology 80, 1475-1494.
Neal, R.M. 2011. MCMC using Hamiltonian dynamics. In: Handbook of Markov Chain Monte Carlo, edited by S. Brooks, A. Gelman, G. Jones, and X.-L. Meng. Chapman & Hall / CRC Press.
These binaries (installable software) and packages are in development.
They may not be fully stable and should be used with caution. We make no claims about them.