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MicropredatorsLimitSignalElaboration_code.R
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MicropredatorsLimitSignalElaboration_code.R
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#########################################################################################################################################
### R code for: "Eavesdropping micropredators as dynamic limiters of sexual signal elaboration and intrasexual competition" ###
### Leavell BC, Beaty LE, McNickle GG, Bernal XE ###
#########################################################################################################################################
########## Procedures ##########
# 1. Assess multicollinearity among variables (GVIF test for collinearity in data)
# 2. Linear Mixed Models; test assumptions
# 3. Generalized Linear Mixed Models; test assumptions
# 4. Linear model selection - cAIC4
# 5. piecewise SEM fit
# 6. Effect Plots
# 7. Deriving Standardized Relevant Ranges
# 8. Alternative piecewiseSEMs
# 8A. Global piecewise SEM without effect of rival males
# 8B. Global piecewise SEM without effect of midges and swats
# 8C. Global piecewise SEM with directionality of path between midges and chucks reversed
# 8D. Global piecewise SEM with directionality of path between midges and chucks reversed and correlated error between Swats and Chucks changed to Chucks ~ Swats
# 9. Investigating a potential case of "Simpson's paradox"
# 9A. Contrasting call rate linear mixed model (LMM) with (original) and without (new) swats
# 9B. Role of ecologically-relevant subgroups in driving potential heterogeneity
# 10. Literature Cited
# Set working directory
setwd("~/")
# Load required libraries
library(ape) # v5.2
library(caper) # v1.0.1
library(piecewiseSEM) # v2.1.0
library(GGally) # v1.4.0
library(lattice) # v0.20.35
library(ggResidpanel) # v0.3.0
library(cAIC4) # v0.9
library(lme4) # v1.1.21
library(MuMIn) # v1.43.6
library(effects) # v4.1.2
library(DiagrammeR) # v1.0.1
library(blmeco) # v1.4
library(DHARMa) # v0.2.4
library(glmmTMB) # v1.0.1
library(bbmle) # v1.0.23.1
library(dplyr) # v1.0.2
# Files needed for script:
# 1. "MicropredatorsLimitSignalElaboration_data.csv"
# 2. "HighstatLibV6.R"
# ("HighstatLibV6.R" is a script for the corvif function (Zuur et al. 2009). It is available online at http://www.highstat.com/book2.htm)
# Read data file
frogdata <- read.csv("~/MicropredatorsLimitSignalElaboration_data.csv")
# Variables
# date = date (day-month-year) of observation
# time = duration (seconds) from beginning of 1st call to beginning of 50th sequential call
# call_rate = # the total number of calls (50), minus one, divided by the time from the beginning of the first call to the beginning of the last call
# chucks = total # of chucks over the 50 sequential calls
# midges = total # of frog-biting midges observed landing on focal frog over 50 sequential calls
# swatcount = total # of swats observed over the 50 sequential call duration
# males_lessthan1m = # of neighbor male competitors present within 1 meter of focal frog
# males_morethan1m = level of perceived abundance of calling conspecifics beyond 1 meter. (0 = only focal frog heard calling, 1 = individual calling frogs could be counted, 2 = calls of frogs overlapping but individuals distinguishable, 3 = full chorus, cannot distinguish individuals). Scale follows Heyer et al. 1994.
## Inspect data
ggpairs(frogdata, cardinality_threshold = 31) #increased cardinality_threshold in order to include date (=30 levels; default only allows for 15 levels)
## Print behavioral data for supplementary figure
ggpairs(frogdata[,-c(1:2)],columnLabels = c("Call rate", "Chucks", "Midges", "Swats", "Males < 1m", "Males > 1m"))
###### 1. Assess multicollinearity among variables #####
collinearity_tab=cbind(frogdata$call_rate,frogdata$chucks,frogdata$midges,frogdata$swatcount,frogdata$males_lessthan1m,frogdata$males_morethan1m)
colnames(collinearity_tab)=c("call_rate","chucks","midges","swatcount","males_lessthan1m","males_morethan1m")
source('~/HighstatLibV6.R', encoding = 'UTF-8') # "HighstatLibV6.R" is a script for the corvif function (Zuur et al. 2009). It is available online at http://www.highstat.com/book2.htm
corvif(collinearity_tab)
#=> all GVIF values are similar and <5 except for males_lessthan1m and males_morethan1m: remove males_morethan1m and try again.
collinearity_tab=cbind(frogdata$call_rate,frogdata$chucks,frogdata$midges,frogdata$swatcount,frogdata$males_lessthan1m)
colnames(collinearity_tab)=c("call_rate","chucks","midges","swatcount","males_lessthan1m")
source('~/HighstatLibV6.R', encoding = 'UTF-8') # "HighstatLibV6.R" is a script for the corvif function (Zuur et al. 2009). It is available online at http://www.highstat.com/book2.htm
corvif(collinearity_tab)
#=> all GVIF values are similar and <5: no indication of collinearity among variables used in the models (Zuur et al. 2007, 2009).
# not using males_morethan1m in models
# Hypothesized pathways for initial piecewiseSEM
# Linear models
# call_rate ~ males_lessthan1m + swatcount + midges
# chucks ~ males_lessthan1m
# midges ~ chucks + males_lessthan1m
# swatcount ~ midges
# Correlated error
# call_rate %~~% chucks
# "date" is random effect for all linear models
# Confirm Initial model meets recommended # of samples/parameter (Grace, Scheiner and Schoolmaster 2015)
# Recommended range is 5 (low) to 20 (plenty)
# 85 samples
# 8 parameters
# 85/8 = approx. 10.6
# Sufficient samples/parameter
####### 2. Linear mixed models #######
# Endogenous: Swatcount
swats_lmm1 <- lmer(swatcount~ midges + (1|date), data=frogdata) # random group intercept
swats_lmm2 <- lmer(swatcount~ midges + (0+midges|date), data=frogdata) # random slope of x w/in group, constant intercept
# FAILED TO CONVERGE swats_lmm3 <- lmer(swatcount~ midges + (1|date) + (0+midges|date), data=frogdata) # uncorrelated random intercept and random slope w/in group
# FAILED TO CONVERGE swats_lmm4 <- lmer(swatcount~ midges + (midges|date), data=frogdata) # random slope of x w/in group, correlated intercept
resid_panel(swats_lmm1) # not good
resid_panel(swats_lmm2) # not good
shapiro.test(resid(swats_lmm1)) # not normal
shapiro.test(resid(swats_lmm2)) # not normal
# Models do not meet assumptions, try GLMM (below)
# Endogenous: Midges
midges_lmm1 <- lmer(midges~ chucks + males_lessthan1m + (1|date), data =frogdata) # random group intercept
# BOUNDARY FIT midges_lmm2 <- lmer(midges~ chucks + males_lessthan1m + (0 + chucks|date), data =frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT midges_lmm3 <- lmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + chucks|date), data =frogdata) # uncorrelated random intercept and random slope w/in group
# BOUNDARY FIT midges_lmm4 <- lmer(midges~ chucks + males_lessthan1m + (chucks|date), data =frogdata) # random slope of x w/in group, correlated intercept
resid_panel(midges_lmm1) # not good
shapiro.test(resid(midges_lmm1)) # not normal
# BOUNDARY FIT midges_lmm5 <- lmer(midges~ chucks + males_lessthan1m + (0 + males_lessthan1m|date), data =frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT midges_lmm6 <- lmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), data =frogdata) # uncorrelated random intercept and random slope w/in group
# BOUNDARY FIT midges_lmm7 <- lmer(midges~ chucks + males_lessthan1m + (males_lessthan1m|date), data =frogdata) # random slope of x w/in group, correlated intercept
# Models do not meet assumptions, try GLMM (below)
# Endogenous: Callrate
callrate_lmm1 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (1|date), data = frogdata) # random group intercept
# NEAR BOUNDARY callrate_lmm2 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (0 + males_lessthan1m|date), data = frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT callrate_lmm3 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (1|date) + (0 + males_lessthan1m|date), data = frogdata) # uncorrelated random intercept and random slope w/in group
# BOUNDARY FIT callrate_lmm4 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (males_lessthan1m|date), data = frogdata) # random slope of x w/in group, correlated intercept
resid_panel(callrate_lmm1) # good
shapiro.test(resid(callrate_lmm1)) # good
# BOUNDARY FIT callrate_lmm5 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (0 + swatcount|date), data = frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT callrate_lmm6 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (1|date) + (0 + swatcount|date), data = frogdata) # uncorrelated random intercept and random slope w/in group
# BOUNDARY FIT callrate_lmm7 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (swatcount|date), data = frogdata) # random slope of x w/in group, correlated intercept
# NEAR BOUNDARY callrate_lmm8 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (0 + midges|date), data = frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT callrate_lmm9 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (1|date) + (0 + midges|date), data = frogdata) # uncorrelated random intercept and random slope w/in group
# FAILED TO CONVERGE callrate_lmm10 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (midges|date), data = frogdata) # random slope of x w/in group, correlated intercept
# callrate_lmm1 is top model; good fit -- keep for SEM
summary(callrate_lmm1)
plot(allEffects(callrate_lmm1))
rsquared(callrate_lmm1)
# Endogenous: Chucks
chucks_lmm1 <- lmer(chucks ~ males_lessthan1m + (1|date), data = frogdata) # random group intercept
chucks_lmm2 <- lmer(chucks ~ males_lessthan1m + (0 + males_lessthan1m|date),data = frogdata) # random slope of x w/in group, constant intercept
# BOUNDARY FIT chucks_lmm3 <- lmer(chucks ~ males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), data = frogdata) # uncorrelated random intercept and random slope w/in group
# FAILED TO CONVERGE chucks_lmm4 <- lmer(chucks ~ males_lessthan1m + (males_lessthan1m|date), data = frogdata) # random slope of x w/in group, correlated intercept
resid_panel(chucks_lmm1) # ok
resid_panel(chucks_lmm2) # ok
shapiro.test(resid(chucks_lmm1)) # good
shapiro.test(resid(chucks_lmm2)) # not great
# Refit from REML to ML for cAIC
chucks_lmm1 <- lmer(chucks ~ males_lessthan1m + (1|date), REML=FALSE, data = frogdata) # random group intercept
chucks_lmm2 <- lmer(chucks ~ males_lessthan1m + (0 + males_lessthan1m|date),REML=FALSE, data = frogdata) # random slope of x w/in group, constant intercept
cAIC(chucks_lmm1)$caic # 838.8899
cAIC(chucks_lmm2)$caic # 860.4867
# top model is chucks_lmm1; fit ok, but compare with GLMM (below)
#refit to REML for overview of summary, plot, rsquared
chucks_lmm1 <- lmer(chucks ~ males_lessthan1m + (1|date), data = frogdata) # random group intercept
summary(chucks_lmm1)
plot(allEffects(chucks_lmm1))
rsquared(chucks_lmm1)
####### 3. Generalized linear mixed models #######
# Endogenous: Swatcount
#sqrt link
swats_glmm1 <- glmer(swatcount~midges+(1|date), family=poisson(link="sqrt"), data=frogdata) # random group intercept
swats_glmm2 <- glmer(swatcount~midges+(0+midges|date), family=poisson(link="sqrt"), data=frogdata) # random slope of x w/in group, constant intercept
swats_glmm3 <- glmer(swatcount~midges+(1|date) + (0+midges|date), family=poisson(link="sqrt"), data=frogdata) # uncorrelated random intercept and random slope w/in group
swats_glmm4 <- glmer(swatcount~midges+(midges|date), family=poisson(link="sqrt"), data=frogdata) # random slope of x w/in group, correlated intercept
resid_panel(swats_glmm1) # good
resid_panel(swats_glmm2) # not good
resid_panel(swats_glmm3) # not awful
resid_panel(swats_glmm4) # not good
shapiro.test(resid(swats_glmm1)) # good
shapiro.test(resid(swats_glmm2)) # not good
shapiro.test(resid(swats_glmm3)) # not good
shapiro.test(resid(swats_glmm4)) # not good
#log link
# LARGE EIGENVALUE swats_glmm5 <- glmer(swatcount~midges+(1|date), family=poisson(link="log"), data=frogdata) # random group intercept
swats_glmm6 <- glmer(swatcount~midges+(0+midges|date), family=poisson(link="log"), data=frogdata) # random slope of x w/in group, constant intercept
swats_glmm7 <- glmer(swatcount~midges+(1|date) + (0+midges|date), family=poisson(link="log"), data=frogdata) # uncorrelated random intercept and random slope w/in group
swats_glmm8 <- glmer(swatcount~midges+(midges|date), family=poisson(link="log"), data=frogdata) # random slope of x w/in group, correlated intercept
resid_panel(swats_glmm6) # not good
resid_panel(swats_glmm7) # ok
resid_panel(swats_glmm8) # not good
shapiro.test(resid(swats_glmm6)) # not good
shapiro.test(resid(swats_glmm7)) # not good
shapiro.test(resid(swats_glmm8)) # not good
#identity link
# ERROR (PIRLS) swats_glmm9 <- glmer(swatcount~midges+(1|date), family=poisson(link="identity"), data=frogdata) # random group intercept
# ERROR (PIRLS) swats_glmm10 <- glmer(swatcount~midges+(0+midges|date), family=poisson(link="identity"), data=frogdata) # random slope of x w/in group, constant intercept
# ERROR (PIRLS) swats_glmm11 <- glmer(swatcount~midges+(1|date) + (0+midges|date), family=poisson(link="identity"), data=frogdata) # uncorrelated random intercept and random slope w/in group
# ERROR (PIRLS) swats_glmm12 <- glmer(swatcount~midges+(midges|date), family=poisson(link="identity"), data=frogdata) # random slope of x w/in group, correlated intercept
cAIC(swats_glmm1) #964.90
cAIC(swats_glmm7) #662.27
# top model is swats_glmm7
summary(swats_glmm7)
plot(allEffects(swats_glmm7)) #unrealistically large values for dependent variable
rsquared(swats_glmm7) # spits out nearly perfect conditional R2... suspect.
#given visual tests of assumptions were not great for swats_glmm7, moving forward with next best model (by cAIC score == swats_glmm1)
summary(swats_glmm1)
plot(allEffects(swats_glmm1))
rsquared(swats_glmm1)
#check for overdispersion w blmeco
dispersion_glmer(swats_glmm1) # indicates overdispersion (value not between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput_swat <- simulateResiduals(fittedModel = swats_glmm1)
plot(simulationOutput_swat) #not great
testDispersion(simulationOutput_swat) #ok
testUniformity(simulationOutput_swat) #ok
testZeroInflation(simulationOutput_swat) #bad
plot(residuals(swats_glmm1)~fitted(swats_glmm1))
# Due to overdispersion from blmeco, adding observational level random effect OLRE to test if overdispersion improves...
# use frog ID as random effect
ID <- c(1:85); frogdata <- cbind(frogdata,ID)
# (Harrison 2014)
swats_glmm1_disper <- glmer(swatcount~midges+(1|date)+(1|ID), family=poisson(link="sqrt"), data=frogdata)
summary(swats_glmm1_disper)
#check dispersion
dispersion_glmer(swats_glmm1_disper) # no evidence of overdispersion (value between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput_swat2 <- simulateResiduals(fittedModel = swats_glmm1_disper)
plot(simulationOutput_swat2) #resid vs predict are bad, looks zero-inflated
plotResiduals(frogdata$midges,simulationOutput_swat2$scaledResiduals)
testDispersion(simulationOutput_swat2) #good
testUniformity(simulationOutput_swat2) #good
##Residual vs predicted lines are also bad for this model when OLRE is added
testZeroInflation(simulationOutput_swat2) #indeed, ZI
#potentially a zero inflated model might resolve resid vs. pred issue, but ZI models are not supported by current version of piecewiseSEM
#Negative binomial?
swats_glmm.nb <- (glmer.nb(swatcount~midges+(1|date), data=frogdata))
dispersion_glmer(swats_glmm.nb) # good
simulationOutput_swat5 <- simulateResiduals(fittedModel = swats_glmm.nb)
plot(simulationOutput_swat5) #resid vs predict are bad, looks zero-inflated
plotResiduals(frogdata$midges,simulationOutput_swat5$scaledResiduals)
testDispersion(simulationOutput_swat5) #good
testUniformity(simulationOutput_swat5) #good
##Residual vs predicted lines are also bad for this model when OLRE is added
testZeroInflation(simulationOutput_swat5) #negative binomial badly zero-inflated, discard
cAIC(swats_glmm1) # cAIC = 964.90
cAIC(swats_glmm1_disper) # cAIC = 748.41
#Choosing swats_glmm1_disper for piecewiseSEM
summary(swats_glmm1_disper)
plot(allEffects(swats_glmm1_disper))
plot(predictorEffect("midges", swats_glmm1_disper), type="response")
rsquared(swats_glmm1_disper)
#Compare swats_glmm1_disper to a hurdle model
# ---- glmmTMB hurdle model ---- #
# -- only midges in zi formula, log link --- #
# truncated_compois(link = "log")
# underdispersed
hurdle1 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_compois(link = "log"))
# truncated_poisson(link = "log")
# underdispersed
hurdle2 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_poisson(link = "log"))
# truncated_nbinom2(link = "log")
#underdispersed
hurdle3 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_nbinom2(link = "log"))
# truncated_nbinom1(link = "log")
# model meets assumptions
# double check res ~ pred
hurdle4 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_nbinom1(link = "log"))
simulationOutput_swat4 <- simulateResiduals(fittedModel = hurdle4)
plot(simulationOutput_swat4)
plotResiduals(frogdata$midges,simulationOutput_swat4$scaledResiduals)
testDispersion(simulationOutput_swat4)
testUniformity(simulationOutput_swat4)
testZeroInflation(simulationOutput_swat4)
# -- midges and (1|date) in zi formula, log link --- #
# truncated_compois(link = "log")
# many warnings
hurdle5 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_compois(link = "log"))
# truncated_poisson(link = "log")
# underdispersed
hurdle6 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_poisson(link = "log"))
# truncated_nbinom2(link = "log")
# model convergence problem
hurdle7 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_nbinom2(link = "log"))
# truncated_nbinom1(link = "log")
# underdispersed
hurdle8 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_nbinom1(link = "log"))
simulationOutput_swat4 <- simulateResiduals(fittedModel = hurdle8)
plot(simulationOutput_swat4)
plotResiduals(frogdata$midges,simulationOutput_swat4$scaledResiduals)
testDispersion(simulationOutput_swat4)
testUniformity(simulationOutput_swat4)
testZeroInflation(simulationOutput_swat4)
# -- midges only in zi formula, sqrt link --- #
# truncated_compois(link = "sqrt")
# took forever, crashed before finishing
#hurdle9 <- glmmTMB(swatcount~midges+(1|date),
# data=frogdata,
# zi=~midges,
# family=truncated_compois(link = "sqrt"))
# truncated_poisson(link = "sqrt")
# underdispersed, KS test bad
hurdle10 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_poisson(link = "sqrt"))
# truncated_nbinom2(link = "sqrt")
# meets assumptions
hurdle11 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_nbinom2(link = "sqrt"))
# truncated_nbinom1(link = "sqrt")
# underdispersed
hurdle12 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~midges,
family=truncated_nbinom1(link = "sqrt"))
simulationOutput_swat4 <- simulateResiduals(fittedModel = hurdle12)
plot(simulationOutput_swat4)
plotResiduals(frogdata$midges,simulationOutput_swat4$scaledResiduals)
testDispersion(simulationOutput_swat4)
testUniformity(simulationOutput_swat4)
testZeroInflation(simulationOutput_swat4)
# -- midges and (1|date) in zi formula, w link = sqrt --- #
# truncated_compois(link = "sqrt")
# took forever, did not finish
# hurdle13 <- glmmTMB(swatcount~midges+(1|date),
# data=frogdata,
# zi=~.,
# family=truncated_compois(link = "sqrt"))
# truncated_poisson(link = "sqrt")
# model convergence problem
#hurdle14 <- glmmTMB(swatcount~midges+(1|date),
# data=frogdata,
# zi=~.,
# family=truncated_poisson(link = "sqrt"))
# truncated_nbinom2(link = "sqrt")
# meets assumptions
hurdle15 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_nbinom2(link = "sqrt"))
# truncated_nbinom1(link = "sqrt")
# underdispersed
hurdle16 <- glmmTMB(swatcount~midges+(1|date),
data=frogdata,
zi=~.,
family=truncated_nbinom1(link = "sqrt"))
simulationOutput_swat4 <- simulateResiduals(fittedModel = hurdle11)
plot(simulationOutput_swat4)
plotResiduals(frogdata$midges,simulationOutput_swat4$scaledResiduals)
testDispersion(simulationOutput_swat4)
testUniformity(simulationOutput_swat4)
testZeroInflation(simulationOutput_swat4)
## -- model selectioin -- ##
# AICtab
AICctab(hurdle1,
hurdle2,
hurdle3,
hurdle4,
hurdle5,
hurdle6,
hurdle7,
hurdle8,
#hurdle9,
hurdle10,
hurdle11,
hurdle12,
#hurdle13,
#hurdle14,
hurdle15,
hurdle16
)
#check dispersion/residuals vs. pred./etc.
simulationOutput_swat4 <- simulateResiduals(fittedModel = hurdle12)
plot(simulationOutput_swat4)
plotResiduals(frogdata$midges,simulationOutput_swat4$scaledResiduals)
testDispersion(simulationOutput_swat4)
testUniformity(simulationOutput_swat4)
testZeroInflation(simulationOutput_swat4)
# dAICc df
# hurdle16 0.0 7 underdispersed
# hurdle12 2.0 6 underdispersed
# hurdle15 2.7 7 ok
# hurdle11 4.7 6 ok
# hurdle5 7.4 7
# hurdle4 7.9 6
# hurdle1 9.3 6
# hurdle7 11.5 7
# hurdle3 13.5 6
# hurdle9 145.1 6
# hurdle10 200.1 5
# hurdle6 224.7 6
# hurdle2 226.8 5
# hurdle8 NA 7
summary(hurdle15) #estimate (sig. negative slope) for midges in count model does not make sense w/ respect to raw data
#reject model
summary(hurdle11) #estimate makes sense (i.e. positive slope)
# meets assumptions
summary(hurdle11) #top model
#estimate and std. error of effect of midges on swatcount very close to the swats_glmm1_disper GLMM in piecewise SEM
# From hurdle model...
# Conditional model:
# Estimate Std. Error z value Pr(>|z|)
# (Intercept) 3.602337 0.393825 9.147 < 2e-16 ***
# midges 0.039266 0.008334 4.712 2.46e-06 ***
# Zero-inflation model:
# Estimate Std. Error z value Pr(>|z|)
# (Intercept) 1.6979 0.8699 1.952 0.05095 .
# midges -0.4155 0.1594 -2.607 0.00912 **
# From GLMM in piecewiseSEM...
# Fixed effects:
# Estimate Std. Error z value Pr(>|z|)
# (Intercept) 3.075105 0.347640 8.846 < 2e-16 ***
# midges 0.037447 0.006006 6.235 4.51e-10 ***
# Comparable effects for non-zero count data between models
#figure
plot(predictorEffects(hurdle11),main = "Swats ~ Midge attacks (Hurdle Model)", ylab = "Swats", xlab = "Midge attacks",
lines=list(col="red"),
rug=FALSE, type = "response")
# Endogenous: Midges
#sqrt link
midges_glmm1 <- glmer(midges~ chucks + males_lessthan1m + (1|date), family = poisson(link="sqrt"), data =frogdata) # random group intercept
midges_glmm2 <- glmer(midges~ chucks + males_lessthan1m + (0 + chucks|date), family = poisson(link="sqrt"), data =frogdata) # random slope of x w/in group, constant intercept
midges_glmm3 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + chucks|date), family = poisson(link="sqrt"), data =frogdata) # uncorrelated random intercept and random slope w/in group
midges_glmm4 <- glmer(midges~ chucks + males_lessthan1m + (chucks|date), family = poisson(link="sqrt"), data =frogdata) # random slope of x w/in group, correlated intercept
midges_glmm5 <- glmer(midges~ chucks + males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="sqrt"), data =frogdata) # random slope of x w/in group, constant intercept
midges_glmm6 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="sqrt"), data =frogdata) # uncorrelated random intercept and random slope w/in group
midges_glmm7 <- glmer(midges~ chucks + males_lessthan1m + (males_lessthan1m|date), family = poisson(link="sqrt"), data =frogdata) # random slope of x w/in group, correlated intercept
resid_panel(midges_glmm1) # good
resid_panel(midges_glmm2) # ok
resid_panel(midges_glmm3) # not good
resid_panel(midges_glmm4) # not good
resid_panel(midges_glmm5) # good
resid_panel(midges_glmm6) # not good
resid_panel(midges_glmm7) # not good
shapiro.test(resid(midges_glmm1)) # good
shapiro.test(resid(midges_glmm2)) # good
shapiro.test(resid(midges_glmm3)) # not good
shapiro.test(resid(midges_glmm4)) # not good
shapiro.test(resid(midges_glmm5)) # good
shapiro.test(resid(midges_glmm6)) # good
shapiro.test(resid(midges_glmm7)) # good
#log link
# LARGE EIGENVALUE midges_glmm8 <- glmer(midges~ chucks + males_lessthan1m + (1|date), family = poisson(link="log"), data =frogdata) # random group intercept
midges_glmm9 <- glmer(midges~ chucks + males_lessthan1m + (0 + chucks|date), family = poisson(link="log"), data =frogdata) # random slope of x w/in group, constant intercept
midges_glmm10 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + chucks|date), family = poisson(link="log"), data =frogdata) # uncorrelated random intercept and random slope w/in group
midges_glmm11 <- glmer(midges~ chucks + males_lessthan1m + (chucks|date), family = poisson(link="log"), data =frogdata) # random slope of x w/in group, correlated intercept
# LARGE EIGENVALUE midges_glmm9 <- glmer(midges~ chucks + males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="log"), data =frogdata) # random slope of x w/in group, constant intercept
# LARGE EIGENVALUE midges_glmm10 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="log"), data =frogdata) # uncorrelated random intercept and random slope w/in group
# LARGE EIGENVALUE midges_glmm11 <- glmer(midges~ chucks + males_lessthan1m + (males_lessthan1m|date), family = poisson(link="log"), data =frogdata) # random slope of x w/in group, correlated intercept
resid_panel(midges_glmm9) # ok
resid_panel(midges_glmm10) # ok
resid_panel(midges_glmm11) # ok
shapiro.test(resid(midges_glmm9)) # good
shapiro.test(resid(midges_glmm10)) # not good
shapiro.test(resid(midges_glmm11)) # not good
#identity link
# PIRLS midges_glmm12 <- glmer(midges~ chucks + males_lessthan1m + (1|date), family = poisson(link="identity"), data =frogdata) # random group intercept
# PIRLS midges_glmm13 <- glmer(midges~ chucks + males_lessthan1m + (0 + chucks|date), family = poisson(link="identity"), data =frogdata) # random slope of x w/in group, constant intercept
# PIRLS midges_glmm14 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + chucks|date), family = poisson(link="identity"), data =frogdata) # uncorrelated random intercept and random slope w/in group
# PIRLS midges_glmm15 <- glmer(midges~ chucks + males_lessthan1m + (chucks|date), family = poisson(link="identity"), data =frogdata) # random slope of x w/in group, correlated intercept
# PIRLS midges_glmm13 <- glmer(midges~ chucks + males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="identity"), data =frogdata) # random slope of x w/in group, constant intercept
# PIRLS midges_glmm14 <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="identity"), data =frogdata) # uncorrelated random intercept and random slope w/in group
# PIRLS midges_glmm15 <- glmer(midges~ chucks + males_lessthan1m + (males_lessthan1m|date), family = poisson(link="identity"), data =frogdata) # random slope of x w/in group, correlated intercept
cAIC(midges_glmm1)$caic # 1713.445
cAIC(midges_glmm2)$caic # Error (PIRLS)
cAIC(midges_glmm5)$caic # Error (PIRLS)
cAIC(midges_glmm9)$caic # 1881.211, but Warnings - failed to converge
cAIC(midges_glmm10)$caic # 1138.573
summary(midges_glmm10)
cAIC(midges_glmm11)$caic # 1166.062, but Warnings - failed to converge
#top model w/o warnings is midges_glmm10
summary(midges_glmm10)
resid_panel(midges_glmm10)
plot(allEffects(midges_glmm10),type="response") #these error bars are enormous...
rsquared(midges_glmm10) # conditional = 1 is suspect
#check for overdispersion w blmeco
dispersion_glmer(midges_glmm10) # indicates overdispersion (value not between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput <- simulateResiduals(fittedModel = midges_glmm10)
plot(simulationOutput) #not great
testDispersion(simulationOutput) #very bad
testUniformity(simulationOutput) #bad
# Due to overdispersion from blmeco, adding observational level random effect OLRE to test if overdispersion improves...
# use frog ID as random effect
# ID <- c(1:85); frogdata <- cbind(frogdata,ID) ... this has already been performed in above code
midges_glmm10_disper <- glmer(midges~ chucks + males_lessthan1m + (1|date) + (0 + chucks|date) + (1|ID), family = poisson(link="log"), data =frogdata) # uncorrelated random intercept and random slope w/in group
summary(midges_glmm10_disper)
#check dispersion
dispersion_glmer(midges_glmm10_disper) # no evidence of overdispersion (value between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput2 <- simulateResiduals(fittedModel = midges_glmm10_disper)
plot(simulationOutput2) #not good
plotResiduals(frogdata$chucks, simulationOutput2$scaledResiduals)
plotResiduals(frogdata$males_lessthan1m, simulationOutput2$scaledResiduals)
testDispersion(simulationOutput2) #not good
testUniformity(simulationOutput2) #not good
#the midges_glmm10_disper is still indicating significant deviation from KS test (p < 0.05)
#reject this model and move to next best model (w/o warnings), which is midges_glmm1
summary(midges_glmm1)
resid_panel(midges_glmm1)
plot(allEffects(midges_glmm1),type="response")
rsquared(midges_glmm1)
#check for overdispersion w blmeco
dispersion_glmer(midges_glmm1) # indicates overdispersion (value not between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput <- simulateResiduals(fittedModel = midges_glmm1)
plot(simulationOutput) #not great
testDispersion(simulationOutput) #ok
testUniformity(simulationOutput) #good
# Due to overdispersion from blmeco, adding observational level random effect OLRE to test if overdispersion improves...
# use frog ID as random effect
# ID <- c(1:85); frogdata <- cbind(frogdata,ID) ... this has already been performed in above code
# (Harrison XA. 2014. Using observation-level random effects to model overdispersion in count data in ecology and evolution. PeerJ 2:e616 https://doi.org/10.7717/peerj.616)
midges_glmm1_disper <- glmer(midges~chucks+males_lessthan1m+(1|date)+(1|ID),family=poisson(link="sqrt"),data=frogdata)
summary(midges_glmm1_disper)
#check dispersion
dispersion_glmer(midges_glmm1_disper) # no evidence of overdispersion (value between 0.75 and 1.4)
#DHARMa check for overdispersion
simulationOutput2 <- simulateResiduals(fittedModel = midges_glmm1_disper)
plot(simulationOutput2) #not great, especially resid vs. pred plot.. could be product of low sample size
# plot by predictors
plotResiduals(frogdata$chucks, simulationOutput2$scaledResiduals)
plotResiduals(frogdata$males_lessthan1m, simulationOutput2$scaledResiduals)
testDispersion(simulationOutput2) #good
testUniformity(simulationOutput2) #good
testZeroInflation(simulationOutput2) #good
#In favor of midge GLMM with OLRE moving forward
resid_panel(midges_glmm1_disper)
shapiro.test(resid(midges_glmm1_disper)) #good
plot(allEffects(midges_glmm1_disper),type="response")
rsquared(midges_glmm1_disper) #predictors do not explain much variance (marginal R2)
# Endogenous: Callrate
# Not fitting GLMMs for callrate due to good fit w/ LMM
# Endogenous: Chucks
#sqrt link
chucks_glmm1 <- glmer(chucks ~ males_lessthan1m + (1|date), family = poisson(link="sqrt"), data = frogdata) # random group intercept
chucks_glmm2 <- glmer(chucks ~ males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="sqrt"), data = frogdata) # random slope of x w/in group, constant intercept
chucks_glmm3 <- glmer(chucks ~ males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="sqrt"), data = frogdata) # uncorrelated random intercept and random slope w/in group
chucks_glmm4 <- glmer(chucks ~ males_lessthan1m + (males_lessthan1m|date), family = poisson(link="sqrt"), data = frogdata) # random slope of x w/in group, correlated intercept
resid_panel(chucks_glmm1) # not good
resid_panel(chucks_glmm2) # not good
resid_panel(chucks_glmm3) # not good
resid_panel(chucks_glmm4) # not good
shapiro.test(resid(chucks_glmm1)) # not good
shapiro.test(resid(chucks_glmm2)) # not good
shapiro.test(resid(chucks_glmm3)) # not good
shapiro.test(resid(chucks_glmm4)) # not good
#log link
chucks_glmm5 <- glmer(chucks ~ males_lessthan1m + (1|date), family = poisson(link="log"), data = frogdata) # random group intercept
chucks_glmm6 <- glmer(chucks ~ males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="log"), data = frogdata) # random slope of x w/in group, constant intercept
chucks_glmm7 <- glmer(chucks ~ males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="log"), data = frogdata) # uncorrelated random intercept and random slope w/in group
chucks_glmm8 <- glmer(chucks ~ males_lessthan1m + (males_lessthan1m|date), family = poisson(link="log"), data = frogdata) # random slope of x w/in group, correlated intercept
resid_panel(chucks_glmm5) # not good
resid_panel(chucks_glmm6) # not good
resid_panel(chucks_glmm7) # not good
resid_panel(chucks_glmm8) # not good
shapiro.test(resid(chucks_glmm5)) # not good
shapiro.test(resid(chucks_glmm6)) # not good
shapiro.test(resid(chucks_glmm7)) # not good
shapiro.test(resid(chucks_glmm8)) # not good
#identity link
# ERROR (PIRLS) chucks_glmm9 <- glmer(chucks ~ males_lessthan1m + (1|date), family = poisson(link="identity"), data = frogdata) # random group intercept
# ERROR (PIRLS) chucks_glmm10 <- glmer(chucks ~ males_lessthan1m + (0 + males_lessthan1m|date), family = poisson(link="identity"), data = frogdata) # random slope of x w/in group, constant intercept
# ERROR (PIRLS) chucks_glmm11 <- glmer(chucks ~ males_lessthan1m + (1|date) + (0 + males_lessthan1m|date), family = poisson(link="identity"), data = frogdata) # uncorrelated random intercept and random slope w/in group
# ERROR (PIRLS) chucks_glmm12 <- glmer(chucks ~ males_lessthan1m + (males_lessthan1m|date), family = poisson(link="identity"), data = frogdata) # random slope of x w/in group, correlated intercept
# no good fitting GLMM, so stick with LMM (above)
####### 4. Linear Model Selection #######
# Endogenous: Swatcount
# LMMs didn't meet assumptions
# Top GLMM is swats_glmm1_disper
swats_glmm1_disper <- glmer(swatcount~midges+(1|date)+(1|ID), family=poisson(link="sqrt"), data=frogdata)
summary(swats_glmm1_disper)
# Endogenous: Midges
# LMMs didn't meet assumptions
# Top GLMM model is midges_glmm1_disper
midges_glmm1_disper <- glmer(midges~chucks+males_lessthan1m+(1|date)+(1|ID),family=poisson(link="sqrt"),data=frogdata)
summary(midges_glmm1_disper)
# Endogenous: Callrate
# Top model is callrate_lmm1
#Make sure refit from ML to REML
callrate_lmm1 <- lmer(call_rate ~ males_lessthan1m + swatcount + midges + (1|date), REML=TRUE, data = frogdata) # random group intercept
summary(callrate_lmm1)
# Endogenous: Chucks
# GLMMs didn't meet assumptions
# Top LMM model is chucks_lmm1
# Refit w/ REML
chucks_lmm1 <- lmer(chucks ~ males_lessthan1m + (1|date), REML=TRUE, data = frogdata) # random group intercept
summary(chucks_lmm1)
####### 5. piecewiseSEM fit #######
pSEM_1 <- psem(
swats_glmm1_disper,
midges_glmm1_disper,
callrate_lmm1,
chucks_lmm1,
chucks %~~% call_rate #We assume chucks and call_rate are correlated
)
summary(pSEM_1) #Fisher's C = 11.041 with P-value = 0.026 and on 4 degrees of freedom
# Conclusion: Reject this pSEM (P < 0.05). Initial causal model does not explain observed data.
# Revise initial model by adding correlated error structure between chucks and swatcount.
pSEM_1.1 <- psem(
swats_glmm1_disper,
midges_glmm1_disper,
callrate_lmm1,
chucks_lmm1,
chucks %~~% call_rate, #We assume chucks and call_rate are correlated
chucks %~~% swatcount # Include this here based on the dSep test in pSEM_1 that revealed a strong relationship between chucks and swatcount
# as we don't have any reason to expect that chucks should drive swatcount, or swats should drive chucks,
#we use correlated errors to account for the correlation
#From https://cran.r-project.org/web/packages/piecewiseSEM/vignettes/piecewiseSEM.html#correlated-errors :
#"Correlated errors reflect the situation where the relationship among the two variables is
#not presumed to be causal and unidirectional,
#but rather that both are being driven by some underlying driver and are therefore appear correlated."
)
summary(pSEM_1.1) #Fisher's C = 0.524 with P-value = 0.769 and on 2 degrees of freedom
summary(pSEM_1.1)$IC
#AIC AICc BIC K n
#38.524 50.378 84.934 19 85
# Model complexity?
# sample size:
summary(pSEM_1.1)$IC$n # sample size = 85
# max nb of paths authorized (Grace et al. 2015):
summary(pSEM_1.1)$IC$n/5 #17 paths authorized
# nb of paths:
nrow(summary(pSEM_1.1)$coefficients) # 9 total paths
# Ratio sample size/paths should be higher than 5:
(summary(pSEM_1.1)$IC$n)/(nrow(summary(pSEM_1.1)$coefficients)) # ratio = 9.44
(summary(pSEM_1.1)$IC$n)/(nrow(summary(pSEM_1.1)$coefficients))>5 # Model complexity is good.
# Conclusion: In accounting for the correlation between chucks and swats, this SEM is an acceptable fit vs. the poor fit of the initial model.
# This model assumes swats and chucks share an underlying driver.
# As chucks and call rate are positively correlated, and swats are a significant negative driver of call rate, it is then not suprising to see the correlation between swats and chucks.
# Also a non-mutually exclusive possibility is that hormones or another factor drive the swat-chuck correlation.
####### 6. Effect Plots #####
#Effect plots w all predictors
plot(allEffects(swats_glmm1_disper), main = "Swats ~ Midge attacks", ylab = "Swats", xlab = "Midge attacks", type="response")
plot(allEffects(midges_glmm1_disper), type="response")
plot(allEffects(callrate_lmm1))
plot(allEffects(chucks_lmm1))
#Single predictor effect plots
#Response: Swats
plot(predictorEffect("midges", swats_glmm1_disper),main = "Swats ~ Midge attacks", ylab = "Swats", xlab = "Midge attacks", rug=FALSE,type="response")
#Response: midges
plot(predictorEffect("chucks", midges_glmm1_disper),main = "Midge attacks ~ Chucks", ylab = "Midge attacks", xlab = "Chucks", rug=FALSE, type="response")
plot(predictorEffect( "males_lessthan1m", midges_glmm1_disper), main = "Midge attacks ~ Males", ylab = "Midge attacks", xlab = "Males", rug=FALSE, type="response")
#Response: call rate
plot(predictorEffect("males_lessthan1m", callrate_lmm1), main = "Call rate ~ Males", ylab = "Call rate", xlab = "Males",rug=FALSE,type="response")
plot(predictorEffect("swatcount", callrate_lmm1), main = "Call rate ~ Swats", ylab = "Call rate", xlab = "Swats",rug=FALSE,type="response")
plot(predictorEffect("midges", callrate_lmm1), main = "Call rate ~ Midges", ylab = "Call rate", xlab = "Midges",rug=FALSE,type="response")
#Response: chucks
plot(predictorEffect("males_lessthan1m", chucks_lmm1), main = "Chucks ~ Males", ylab = "Chucks", xlab = "Males",rug=FALSE,type="response")
# Visual comparison of swats_glmm1_disper model with hurdle model
# hurdle figure
plot(predictorEffects(hurdle11),main = FALSE, ylab = "Swats", xlab = "Midge attacks", ylim=c(0,140),
lines=list(col="red"),
rug=FALSE, type = "response")
# swats_glmm1_disper
plot(predictorEffect("midges", swats_glmm1_disper),main = FALSE, ylab = "Swats", xlab = "Midge attacks", ylim=c(0,140),
rug=FALSE,type="response", add=TRUE)
#########################################
### QUERIES TO OBTAIN STANDARDIZED COEFS
# For a reference, see GRACE AND BOLLEN 2005. BULLETIN OF THE ECOLOGICAL SOCIETY OF AMERICA.
#########################################
# Get standardized coefficients by using "relative range" method outlined in
# (Grace and Bollen 2005)
# code from (Grace et al. 2012)
####### EXAMPLE from Grace et al. 2012 Ecosphere ########################################################
### FOR NATR ~ SURR + WET ###############################################
# log(natr.hat[i]) <- b7.0 +b7.1*wet[i] +b7.2*surr[i]
# b7.0= 3.9928
# b7.1= -0.0024
# b7.2= -0.1646
# range(natr) # 62 - 11= 51
#
# natr.predict.1 <- b7.0 + b7.1*min(wet) +b7.2*mean(surr); print(natr.predict.1) # Scenario1: wet at min and surr at mean
# natr.predict.2 <- b7.0 + b7.1*max(wet) +b7.2*mean(surr); print(natr.predict.2) # Scenario2: wet at max and surr at mean
# natr.predict.3 <- b7.0 + b7.1*mean(wet) +b7.2*min(surr); print(natr.predict.3) # Scenario3: wet at mean and surr at min
# natr.predict.4 <- b7.0 + b7.1*mean(wet) +b7.2*max(surr); print(natr.predict.4) # Scenario4: wet at mean and surr at max
#
# b7.1.std <- (exp(natr.predict.2)-exp(natr.predict.1))*(1/51); print(b7.1.std)
# b7.2.std <- (exp(natr.predict.4)-exp(natr.predict.3))*(1/51); print(b7.2.std)
##########################################################################################################
####### 7. Deriving Standardized Relevant Ranges for Global piecewiseSEM ######
# We chose to use the empirical ranges as the relevant ranges
range(frogdata$swatcount) # == min: 0; max: 90
range(frogdata$midges) # == min: 0; max: 153
range(frogdata$males_lessthan1m) # == min: 0; max: 4
range(frogdata$chucks) # == min: 0; max: 146
range(frogdata$call_rate) # == min: 0.13; max: 0.63
## 1. FOR swatcount ~ midges + (1|date), family=poisson(link="sqrt") ################### This is a GLMM
# sqrt(swatcount.hat[i]) ~ b1.0 + b1.1*midges[i]
b1.0 = fixef(swats_glmm1_disper)[1] # 3.075105 intercept
b1.1 = fixef(swats_glmm1_disper)[2] # 0.03744661 midges
range(frogdata$swatcount) # 90 - 0 = 90
swats.predict.1 <- b1.0 + b1.1*min(frogdata$midges) # Scenario1: midges at min
swats.predict.2 <- b1.0 + b1.1*max(frogdata$midges) # Scenario2: midges at max
b1.1.std <- ((swats.predict.2)^2-(swats.predict.1)^2)*(1/90); print(b1.1.std) # Std. Estimate = 0.7562425 (swats <- midges)
## 2. FOR midges ~ chucks + males_lessthan1m + (1|date), family = poisson(link="sqrt") ################### This is a GLMM
# sqrt(midges.hat[i]) ~ b2.0 + b2.1*chucks[i] + b2.2*males_lessthan1m[i]
b2.0 = fixef(midges_glmm1_disper)[1] # 6.229556 intercept
b2.1 = fixef(midges_glmm1_disper)[2] # -0.01369288 chucks
b2.2 = fixef(midges_glmm1_disper)[3] # 0.09076195 males_lessthan1m
range(frogdata$midges) # 153 - 0 = 153
midges.predict.1 <- b2.0 + b2.1*min(frogdata$chucks) +b2.2*mean(frogdata$males_lessthan1m); print(midges.predict.1) # Scenario1: chucks at min and males_lessthan1m at mean
midges.predict.2 <- b2.0 + b2.1*max(frogdata$chucks) +b2.2*mean(frogdata$males_lessthan1m); print(midges.predict.2) # Scenario2: chucks at max and males_lessthan1m at mean
midges.predict.3 <- b2.0 + b2.1*mean(frogdata$chucks) +b2.2*min(frogdata$males_lessthan1m); print(midges.predict.3) # Scenario3: chucks at mean and males_lessthan1m at min
midges.predict.4 <- b2.0 + b2.1*mean(frogdata$chucks) +b2.2*max(frogdata$males_lessthan1m); print(midges.predict.4) # Scenario4: chucks at mean and males_lessthan1m at max
b2.1.std <- ((midges.predict.2)^2-(midges.predict.1)^2)*(1/153); print(b2.1.std) # Std. Estimate = -0.1440966 (midges <- chucks)
b2.2.std <- ((midges.predict.4)^2-(midges.predict.3)^2)*(1/153); print(b2.2.std) # Std. Estimate = 0.02591771 (midges <- males_lessthan1m)
## 3. FOR call_rate ~ males_lessthan1m + swatcount + midges + (1|date) ################### This is a LMM
# call_rate.hat[i] ~ b4.0 + b4.1*males_lessthan1m[i]
b3.0 = fixef(callrate_lmm1)[1] # 0.4511456 (intercept)
b3.1 = fixef(callrate_lmm1)[2] # 0.009765517 (males_lessthan1m)
b3.2 = fixef(callrate_lmm1)[3] # -0.00166919 (swatcount)
b3.3 = fixef(callrate_lmm1)[4] # -0.0002899801 (midges)
range(frogdata$call_rate) # 0.63 - 0.13 = 0.50
callrate.predict.1 <- b3.0 + b3.1*min(frogdata$males_lessthan1m) + b3.2*mean(frogdata$swatcount) + b3.3*mean(frogdata$midges); print(callrate.predict.1) # Scenario1: males at min
callrate.predict.2 <- b3.0 + b3.1*max(frogdata$males_lessthan1m) + b3.2*mean(frogdata$swatcount) + b3.3*mean(frogdata$midges); print(callrate.predict.2) # Scenario2: males at max
callrate.predict.3 <- b3.0 + b3.1*mean(frogdata$males_lessthan1m) + b3.2*min(frogdata$swatcount) + b3.3*mean(frogdata$midges); print(callrate.predict.3) # Scenario3: swatcount at min
callrate.predict.4 <- b3.0 + b3.1*mean(frogdata$males_lessthan1m) + b3.2*max(frogdata$swatcount) + b3.3*mean(frogdata$midges); print(callrate.predict.4) # Scenario4: swatcount at max
callrate.predict.5 <- b3.0 + b3.1*mean(frogdata$males_lessthan1m) + b3.2*mean(frogdata$swatcount) + b3.3*min(frogdata$midges); print(callrate.predict.5) # Scenario5: midges at min
callrate.predict.6 <- b3.0 + b3.1*mean(frogdata$males_lessthan1m) + b3.2*mean(frogdata$swatcount) + b3.3*max(frogdata$midges); print(callrate.predict.6) # Scenario6: midges at max
b3.1.std <- ((callrate.predict.2)-(callrate.predict.1))*(1/0.5); print(b3.1.std) #Std. Estimate = 0.07812414 (callrate <- males_lessthan1m)
b3.2.std <- ((callrate.predict.4)-(callrate.predict.3))*(1/0.5); print(b3.2.std) #Std. Estimate = -0.3004542 (callrate <- swatcount)
b3.3.std <- ((callrate.predict.6)-(callrate.predict.5))*(1/0.5); print(b3.3.std) #Std. Estimate = -0.0887339 (callrate <- midges)
## 4. FOR chucks ~ males_lessthan1m + (1|date) ################### This is a LMM
# chucks.hat[i] ~ b4.0 + b4.1*males_lessthan1m[i]
b4.0 = fixef(chucks_lmm1)[1] # 44.14221
b4.1 = fixef(chucks_lmm1)[2] # 8.057453
range(frogdata$chucks) # 146 - 0 = 146
chucks.predict.1 <- b4.0 + b4.1*min(frogdata$males_lessthan1m); print(chucks.predict.1) # Scenario1: males at min
chucks.predict.2 <- b4.0 + b4.1*max(frogdata$males_lessthan1m); print(chucks.predict.2) # Scenario2: males at max
b4.1.std <- ((chucks.predict.2)-(chucks.predict.1))*(1/146); print(b4.1.std) #Std. Estimate = 0.2207521 (chucks <- males_lessthan1m)
####### Summary: Standardized Estimates (Relative Ranges Method) for Global piecewiseSEM ##########
#### DIRECT EFFECTS ####
# swats <- midges Std. Estimate = 0.756 (0.7562425)
# midges <- chucks Std. Estimate = -0.144 (-0.1440966)
# midges <- males_lessthan1m Std. Estimate = 0.026 (0.02591771)
# callrate <- males_lessthan1m Std. Estimate = 0.078 (0.07812414)
# callrate <- swatcount Std. Estimate = -0.300 (-0.3004542)
# callrate <- midges Std. Estimate = -0.089 (-0.0887339)
# chucks <- males_lessthan1m Std. Estimate = 0.221 (0.2207521)
#### INDIRECT EFFECTS ####
# Rival males to Midges (via Chucks)
# (Rival males to Chucks) * (Chucks to Midges)
0.2207521*-0.1440966 # = -0.032
# Rival males to Call rate (via Midges)
# (Rival males to Midges) * (Midges to Call rate)
0.02591771*-0.0887339 # = -0.002
# Rival males to Call rate (via Midges & Swats)
# (Rival males to Midges) * (Midges to Swats) * (Swats to Call rate)
0.02591771*0.7562425*-0.3004542 # = -0.006
# Rival males to Call rate (via Chucks & Midges)
# (Rival males to Chucks) * (Chucks to Midges) * (Midges to Call rate)
0.2207521*-0.1440966*-0.0887339 # = 0.003
# Rival males to Call rate (via Chucks & Midges & Swats)
# (Rival males to Chucks) * (Chucks to Midges) * (Midges to Swats) * (Swats to Call rate)
0.2207521*-0.1440966*0.7562425*-0.3004542 # = 0.007
# Midges to Call rate (via Swats)
# (Midges to Swats) * (Swats to Call rate)
0.7562425*-0.3004542 # = -0.227
#### TOTAL EFFECTS ####
# Rival males to Midges
# (Rival males to Chucks) * (Chucks to Midges) + (Rival males to Midges)
0.2207521*-0.1440966+0.02591771 # = -0.006 (-0.005891917)
# Rival males to Call rate
# [(Total Effect: Rival males to Midges) * (Midges to Swats) * ( Swats to Call rate)] + [(Total Effect: Rival males to Midges) * (Midges to Call rate)] + (Rival males to Call rate)
(-0.005891917*0.7562425*-0.3004542) + (-0.005891917*-0.0887339)+0.07812414 # = 0.080 (0.07998569)
# Midges to Call rate
# (Midges to Swats) * ( Swats to Call rate) + (Midges to Call rate)
0.7562425*-0.3004542+-0.0887339 # = -0.316
##############################################################################
####### 8. Alternative piecewiseSEMs ########
####### 8A. Global piecewise SEM without effect of rival males ######
#i.
# Males are not a part of swat model
nomale_midges_glmm1_disper <- glmer(midges ~ chucks + (1 | date) + (1 | ID), family=poisson("sqrt"),data=frogdata)
nomale_callrate_lmm1 <- lmer(call_rate ~ swatcount + midges + (1 | date), data=frogdata)
# There will be no chucks model because the only predictor is males
#ii.
# Check model assumptions
summary(nomale_midges_glmm1_disper)
resid_panel(nomale_midges_glmm1_disper)
dispersion_glmer(nomale_midges_glmm1_disper) #No overdispersion
#DHARMa check for overdispersion
simulationOutput_midges1 <- simulateResiduals(fittedModel = nomale_midges_glmm1_disper)
plot(simulationOutput_midges1) #ok
testDispersion(simulationOutput_midges1) #good
testUniformity(simulationOutput_midges1) #good
summary(nomale_callrate_lmm1)
resid_panel(nomale_callrate_lmm1)
# Now substitute these models into the SEM
pSEM_2 <- psem(
swats_glmm1_disper,
nomale_midges_glmm1_disper,
nomale_callrate_lmm1,
#chucks_lmm1 no chucks model because the only predictor is males
frogdata$males_lessthan1m~1, #In order for AIC values of Nested SEMs to be compared, all variables must be present. Use "males_lessthan1m~1" to include males in the d-sep test to make comparison fair.
#This follows instructions by Jarrett Byrnes (https://github.com/jebyrnes/semclass/blob/master/lecture_pdfs/Local_Estimation.pdf)
#And http://jslefche.github.io/piecewiseSEM/articles/piecewiseSEM.html
chucks %~~% call_rate, #We assume chucks and call_rate are correlated
chucks %~~% swatcount # Include this here based on the dSep test in pSEM_1 that revealed a strong relationship between chucks and swatcount
# as we don't have any reason to expect that chucks should drive swatcount,
#we use correlated errors to account for the correlation
#From https://cran.r-project.org/web/packages/piecewiseSEM/vignettes/piecewiseSEM.html#correlated-errors :
#"Correlated errors reflect the situation where the relationship among the two variables is
#not presumed to be causal and unidirectional,
#but rather that both are being driven by some underlying driver and are therefore appear correlated."
)
summary(pSEM_2) # Fisher's C = 3.252 with P-value = 0.777 and on 6 degrees of freedom
AIC(pSEM_1.1, aicc=TRUE) #50.378
AIC(pSEM_2, aicc=TRUE) #35.02
AIC(pSEM_1.1, aicc=TRUE) - AIC(pSEM_2, aicc=TRUE) # = 15.358 top model is pSEM_2
#Conclusion: Removing males from the SEM results in an acceptable, and indeed better, model fit than pSEM_1.1.
# Midges (and thus swats) are important drivers of call dynamics in túngara frogs.
####### Deriving Standardized Relevant Ranges for Global piecewise SEM without effect of rival males ######
# We chose to use the empirical ranges as the relevant ranges
range(frogdata$swatcount) # == min: 0; max: 90
range(frogdata$midges) # == min: 0; max: 153
range(frogdata$males_lessthan1m) # == min: 0; max: 4
range(frogdata$chucks) # == min: 0; max: 146
range(frogdata$call_rate) # == min: 0.13; max: 0.63