Introduction

Gene expression changes contribute to complex trait variations in both individuals and populations. However, how gene expression influences changes of complex traits over macroevolutionary timescales remains poorly understood. Snake venoms are proteinaceous cocktails where the expression of each toxin can be quantified and mapped to a distinct genomic locus and traced for millions of years. Using a phylogenetic generalized linear mixed model, we analysed expression data of toxin genes from 52 snake species spanning the three venomous snake families, and estimated phylogenetic covariance, which acts as a measure of evolutionary constraint. We find that evolution of toxin combinations is not constrained. However, while all combinations are in principle possible, the actual dimensionality of phylomorphic space is low, with envenomation strategies focused around only four major toxins: metalloproteases, three-finger toxins, serine proteases, and phospholipases A2. While most extant snakes prioritize either a single or a combination of major toxins, they are repeatedly recruited and lost. We find that over macroevolutionary timescales the venom phenotypes were not shaped by phylogenetic constraints, which include important microevolutionary constraints such as epistasis and pleiotropy, but more likely by ecological filtering that permits a few optimal solutions. As a result, phenotypic optima were repeatedly attained by distantly related species. These results indicate that venoms evolve by selection on biochemistry of prey envenomation, which permit diversity though parallelism and impose strong limits, since only a few of the theoretically possible strategies seem to work well and are observed in extant snakes.

library(phytools) #for ASR
library(MCMCglmm) # Estimating PCOV 
library(ggplot2) # plotting
library(reshape2) # data formatting
library(ggrepel) # plotting
library(scatterpie) # pie charts on PCA
library(factoextra) # data formatting
library(surface) #Convergence analysis
library(knitr) #rmd
library(kableExtra) #rmd
library(ade4)# Mantel test

options(knitr.table.format = "html")

Phylogenetic covariance analysis.

#Tree
snphylo = read.nexus("/Users/agneesh-barua/Dropbox (OIST)/R Training/VC_project/Final_tree.nex")
is.ultrametric(snphylo)
## [1] TRUE
#data
phydata = read.csv("/Users/agneesh-barua/Dropbox (OIST)/R Training/VC_project/Data csv/New data/StvNewdata.csv", header = TRUE)
row.names(phydata) <- phydata$species

Supplementary Table 1: Gene expression values and references

species TFTx BPP.CNP CRISP CTL GF KSPI LAAO PLA2 SVMP SVSP Total Family Author Tech Ref
Boiga_irregularis 0.67475 0.01400 0.03835 0.02271 0.00025 0.00100 0.00000 0.00040 0.24854 0.00000 1.00000 Colubridae McGivern et al. (2014) Illumina https://doi.org/10.1186/1471-2164-15-1061
Dispholidus_typus 0.06200 0.00000 0.05400 0.00400 0.00100 0.00150 0.00000 0.07600 0.74640 0.05500 0.99990 Colubridae Pla et al. (2017) Illumina https://doi.org/10.1016/j.bbagen.2017.01.020
Hypsiglena_torquata 0.00070 0.07359 0.16684 0.06176 0.00012 0.00056 0.00000 0.00000 0.68666 0.00000 0.99023 Colubridae McGivern et al. (2014) Illumina https://doi.org/10.1186/1471-2164-15-1061
Opheodrys_aestivus 0.13300 0.00000 0.00000 0.00000 0.00000 0.00000 0.00000 0.03600 0.00000 0.00000 0.16900 Colubridae Hargreaves et al. (2014) Illumina https://doi.org/10.1016/j.toxicon.2014.10.004
Pantherophis_guttatus 0.01000 0.00000 0.02200 0.00000 0.00000 0.00000 0.00000 0.12900 0.00000 0.00000 0.16100 Colubridae Hargreaves et al. (2014) Illumina https://doi.org/10.1016/j.toxicon.2014.10.004
Phalotris_mertensi 0.01800 0.00140 0.00330 0.21600 0.00040 0.43900 0.02300 0.00000 0.22040 0.00220 0.92370 Colubridae Campos et al. (2016) ABI 3100 https://doi.org/10.1093/gbe/evw149
Philodryas_olfersii 0.00000 0.22400 0.10400 0.05800 0.00000 0.00000 0.00000 0.00000 0.52400 0.08900 0.99900 Colubridae Ching et al. (2006) ABI 3100 https://doi.org/10.1016/j.febslet.2006.07.010
Thamnodynastes_strigatus 0.00200 0.00000 0.01900 0.05400 0.00000 0.00000 0.00000 0.00000 0.84400 0.00700 0.92600 Colubridae Ching et al. (2011) ABI 3100 https://dx.doi.org/10.1021/pr200876c
Bungarus_flaviceps 0.39290 0.00000 0.00440 0.00440 0.00000 0.24060 0.00000 0.35320 0.00000 0.00000 0.99550 Elapidae Siang et al. (2010) ABI PRISM https://dx.doi.org/10.1186/1471-2199-11-24
Bungarus_multicinctus 0.64700 0.00000 0.00700 0.03800 0.00200 0.10600 0.00100 0.17400 0.00000 0.00100 0.97600 Elapidae Jiang et al. (2011) ABI 3100 https://doi.org/10.1186/1471-2164-12-1
Drysdalia_coronoides 0.86000 0.00000 0.02800 0.00000 0.01400 0.00000 0.00000 0.00240 0.00000 0.09200 0.99640 Elapidae Chatrath et al. (2011) ABI PRISM https://dx.doi.org/10.1021/pr1008916
Micrurus_altirostris 0.86400 0.00000 0.00000 0.01300 0.00000 0.03800 0.00000 0.04900 0.00600 0.00000 0.97000 Elapidae Correa-Netto et al. (2011) ABI 3100 https://doi.org/10.1016/j.jprot.2011.04.003
Micrurus_corallinus 0.20300 0.29530 0.00000 0.06700 0.01830 0.00150 0.00840 0.37680 0.02100 0.00100 0.99230 Elapidae Aird et al. (2017) Illumina https://dx.doi.org/10.3390/toxins9060187
Micrurus_fulvius 0.25160 0.00000 0.00000 0.00799 0.01600 0.02154 0.00880 0.64947 0.02947 0.00000 0.98487 Elapidae Margres et al. (2013) Illumina https://doi.org/10.1186/1471-2164-14-531
Micrurus_lemniscatus 0.67220 0.00200 0.00240 0.00480 0.01420 0.07150 0.00730 0.17200 0.01120 0.00100 0.95860 Elapidae Aird et al. (2017) Illumina https://dx.doi.org/10.3390/toxins9060187
Micrurus_spixii 0.62640 0.00000 0.00000 0.00000 0.00000 0.00000 0.00000 0.36390 0.00000 0.00000 0.99030 Elapidae Aird et al. (2017) Illumina https://dx.doi.org/10.3390/toxins9060187
Micrurus_surinamensis 0.60240 0.02670 0.00000 0.00340 0.07880 0.01070 0.00000 0.26230 0.00140 0.00000 0.98570 Elapidae Aird et al. (2017) Illumina https://dx.doi.org/10.3390/toxins9060187
Naja_atra 0.95800 0.00000 0.00700 0.00600 0.00100 0.00100 0.00100 0.01200 0.00400 0.00000 0.99000 Elapidae Jiang et al. (2011) ABI 3100 https://doi.org/10.1186/1471-2164-12-1
Naja_kaouthia 0.82000 0.00000 0.00000 0.00500 0.00300 0.00100 0.00200 0.13600 0.01100 0.00000 0.97800 Elapidae Xu et al. (2017) Illumina https://doi.org/10.1016/j.jprot.2017.02.018
Ophiophagus_hannah 0.43000 0.00000 0.08700 0.00000 0.00800 0.01000 0.05700 0.04000 0.24400 0.00700 0.88300 Elapidae Tan et al. (2015) Illumina https://dx.doi.org/10.1186/s12864-015-1828-2
Pseudonaja_textilis 0.10633 0.00000 0.03846 0.00000 0.00452 0.01810 0.00000 0.18778 0.00000 0.42534 0.78053 Elapidae Reeks et al. (2016) Roche GS FLX https://doi.org/10.1016/j.jprot.2015.11.019
Agkistrodon_piscivorus 0.00000 0.04570 0.00000 0.06080 0.00000 0.00000 0.00000 0.38960 0.34900 0.08300 0.92810 Viperidae Aronow (2018) Illumina https://search.proquest.com/openview/2997dede40d17e51e7dabfd03d9762df/1?pq-origsite=gscholar&cbl=18750&diss=y
Atropoides_picadoi 0.00400 0.16400 0.02600 0.04800 0.00600 0.00010 0.03100 0.02300 0.56100 0.10900 0.97210 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Bothriechis_lateralis 0.01600 0.23600 0.03200 0.03100 0.01800 0.00020 0.03400 0.03000 0.44900 0.10400 0.95020 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Bothriechis_schlegelii 0.03600 0.15300 0.03600 0.00200 0.04900 0.00120 0.03200 0.09200 0.26300 0.20500 0.86920 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Bothrops_alternatus 0.00000 0.11800 0.02200 0.16500 0.02500 0.00000 0.01800 0.01400 0.58500 0.05100 0.99800 Viperidae Pereira de Paula et al. (2014) MegaBACE (GE) https://doi.org/10.1016/j.cbd.2014.09.001
Bothrops_asper 0.00800 0.15000 0.00800 0.01000 0.00000 0.00030 0.04200 0.17800 0.41600 0.13400 0.94630 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Bothrops_atrox 0.00000 0.07100 0.00500 0.04800 0.01000 0.00000 0.01400 0.14600 0.61600 0.08200 0.99200 Viperidae Neiva et al. (2009) MegaBACE (GE) https://doi.org/10.1016/j.toxicon.2009.01.006
Bothrops_colombiensis 0.00000 0.01000 0.02000 0.04000 0.02000 0.00000 0.02000 0.30000 0.37000 0.12000 0.90000 Viperidae Suntravat et al. (2016) ABI 3730 https://doi.org/10.1186/s12867-016-0059-7
Bothrops_insularis 0.00000 0.15800 0.01500 0.14200 0.05100 0.00000 0.03500 0.05400 0.43200 0.11200 0.99900 Viperidae Valente et al. (2009) ABI 3100 https://doi.org/10.1016/j.jprot.2009.01.001
Bothrops_jararaca 0.00000 0.04250 0.00400 0.13650 0.02900 0.00200 0.04050 0.03600 0.43600 0.26100 0.98750 Viperidae Goncalves-Machado et al. (2016) Roche GS FLX https://doi.org/10.1016/j.jprot.2015.04.029
Bothrops_jararacussu 0.00000 0.01695 0.03390 0.08475 0.00000 0.00000 0.01695 0.56290 0.25820 0.01695 0.99060 Viperidae Kashima et al. (2004) ABI 3100 https://doi.org/10.1016/j.biochi.2004.02.002
Bothrops_moojeni 0.00060 0.00950 0.00570 0.02670 0.00000 0.00440 0.01270 0.09020 0.46630 0.26180 0.87790 Viperidae Amorim et al. (2017) Illumina https://doi.org/10.1016/j.toxicon.2017.10.025
Bothrops_neuwiedi 0.00000 0.17400 0.00700 0.08600 0.00360 0.00000 0.02900 0.26400 0.36000 0.05300 0.97660 Viperidae Rodigues et al. (2012) MegaBACE (GE) https://doi.org/10.1016/j.jprot.2012.03.028
Cerrophidion_godmani 0.00600 0.08300 0.02600 0.00200 0.00600 0.00000 0.05500 0.07900 0.42200 0.28100 0.96000 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Crotalus_adamanteus 0.00000 0.00726 0.01317 0.22155 0.00023 0.00054 0.05309 0.07810 0.24410 0.19979 0.81783 Viperidae Rokyta et al. (2012) Illumina https://doi.org/10.1186/1471-2164-13-312
Crotalus_durissus 0.00000 0.00000 0.00000 0.00000 0.04000 0.00000 0.00000 0.88000 0.03000 0.02500 0.97500 Viperidae Boldrini-Franca et al. (2009) MegaBACE (GE) https://doi.org/10.1016/j.biochi.2009.02.001
Crotalus_horridus 0.00000 0.12519 0.00775 0.00213 0.02285 0.00119 0.01106 0.22772 0.00111 0.58229 0.98129 Viperidae Rokyta et al. (2013) Illumina https://doi.org/10.1186/1471-2164-14-394
Crotalus_simus 0.12700 0.13400 0.00000 0.02400 0.00000 0.00150 0.03800 0.12100 0.06500 0.28100 0.79150 Viperidae Durban et al. (2011) Roche GS FLX https://doi.org/10.1186/1471-2164-12-259
Crotalus_scutulatus 0.00000 0.20121 0.00481 0.16943 0.02000 0.00021 0.00925 0.18171 0.14363 0.11692 0.84717 Viperidae Strickland et al. (2018) Illumina https://dx.doi.org/10.3390/toxins10040135
Gloydius_intermedius 0.00000 0.04900 0.04900 0.04900 0.14600 0.00000 0.02400 0.12200 0.17000 0.22000 0.82900 Viperidae Yang et al. (2015) MegaBACE (GE) https://doi.org/10.1016/j.toxicon.2015.08.010
Lachesis_muta 0.01600 0.73200 0.00000 0.05800 0.00300 0.00000 0.03700 0.04700 0.05900 0.03500 0.98700 Viperidae Lnacio et al. (2007) ABI 3100 https://doi.org/10.1534/genetics.106.056515
Ovophis_okinavensis 0.00000 0.00000 0.00000 0.00579 0.00008 0.00000 0.00623 0.00647 0.04220 0.86176 0.92253 Viperidae Aird et al. (2013) Illumina https://doi.org/10.1186/1471-2164-14-790
Protobothrops_elegans 0.00000 0.01400 0.00001 0.00200 0.00600 0.00000 0.00500 0.77100 0.08100 0.10400 0.98301 Viperidae Aird et al. (2015) Illumina https://doi.org/10.1186/s12864-015-1832-6
Protobothrops_mucrosquamatus 0.00000 0.00000 0.01018 0.01018 0.00647 0.00000 0.01200 0.25746 0.39966 0.28076 0.97671 Viperidae Aird et al. (2017) Illumina https://doi.org/10.1093/gbe/evx199
Protobothrops_flavoviridis 0.00000 0.00000 0.03854 0.15356 0.00960 0.00000 0.09071 0.32136 0.26975 0.11054 0.99406 Viperidae Aird et al. (2013) Illumina https://doi.org/10.1186/1471-2164-14-790
Sistrurus_catenatus 0.00830 0.00280 0.06670 0.01390 0.06670 0.00000 0.03610 0.28060 0.12220 0.37500 0.97230 Viperidae Pahari et al. (2007) ABI 3100 https://doi.org/10.1186/1471-2199-8-115
Bitis_gabonica 0.00000 0.01813 0.00000 0.07591 0.03626 0.09064 0.05665 0.05665 0.37729 0.14729 0.85882 Viperidae Francischetti et al. (2004) Beckman Coulter https://doi.org/10.1016/j.gene.2004.03.024
Echis_carinatus 0.00000 0.00000 0.01934 0.17602 0.00580 0.00193 0.01161 0.08317 0.64603 0.04836 0.99226 Viperidae Casewell et al. (2009) ABI 3730 https://doi.org/10.1186/1471-2164-10-564
Echis_coloratus 0.00000 0.00000 0.05426 0.10543 0.00620 0.00000 0.04031 0.05426 0.68527 0.04496 0.99069 Viperidae Casewell et al. (2009) ABI 3730 https://doi.org/10.1186/1471-2164-10-564
Echis_ocellatus 0.00000 0.00000 0.00297 0.10979 0.02671 0.00193 0.01187 0.08605 0.73591 0.02374 0.99897 Viperidae Casewell et al. (2009) ABI 3730 https://doi.org/10.1186/1471-2164-10-564
Echis_pyramidum 0.00000 0.00000 0.00000 0.24011 0.00582 0.00000 0.02639 0.20580 0.50000 0.01979 0.99791 Viperidae Casewell et al. (2009) ABI 3730 https://doi.org/10.1186/1471-2164-10-564 
#normalize libraries 
phydata.norm <- phydata
phydata.norm[,2:11] <- t(scale(t(phydata.norm[,2:11])))
phydata.norm$Sum.of.10<-NULL
phydata.norm$Others<-NULL
phydata.norm$Family <-NULL
phydata.norm$Total<-NULL
phydata.norm$Author<-NULL
phydata.norm$Ref<-NULL

The goal of the generalised linear mixed model is to model the effect of phylogeny on the 10 components of snake venom. All 10 snake venom toxins were normalized and passed as multi-response vectors by forming a matrix using the cbind function in R. trait was used to fit different response values, allowing them to have different means. The random effects were added as us(traits):species which allows us to obtain covariance across trait that are influenced by the effect of species(phylogeny), accounting for the covariance between two response variables. In our case where we have used phylogeny as the random effect, the vectors of random effects are associated with the inverse relationship matrix that we obtain via the inv.phylo command. The MCMC was run for a total of 20 million iterations, with burnin and thinning values of 1 million and 1,500 respectively.

inv.phylo = inverseA(snphylo, nodes = "TIPS", scale =TRUE)

priors1 <- list(G=list(G1=list(V=diag(10), nu=0.002)), R=list(V=diag(10), nu= 0.002)) # standard prior
ptm <- proc.time()
cov_model.1<-MCMCglmm(cbind(TFTx,BPP.CNP,CRISP,CTL,GF,KSPI,LAAO,PLA2,SVMP,SVSP) ~ trait:1, 
                    random = ~us(trait):species,
                    rcov = ~us(trait):units,
                    family=rep("gaussian",10), 
                    ginverse=list(species=inv.phylo$Ainv), 
                    prior=priors1,
                    data=phydata.norm[,1:12], 
                    nitt=20000000, burnin=1000000, thin=15, pr = TRUE) #20million, 1million burn-in
saveRDS(cov_model.1, "cov_model_OnSango_sprandom_Final.nosize.20mil.rds")

Diagnostics

cov_model.1<-readRDS("/Users/agneesh-barua/Dropbox (OIST)/R Training/VC_project/GLMM/OnSango/Final/Post_Nov/cov_model_OnSango_sprandom_Final.nosize.20mil.rds")

Covariance model summary

summary(cov_model.1)
## 
##  Iterations = 1000001:19999001
##  Thinning interval  = 1500
##  Sample size  = 12667 
## 
##  DIC: -1520.484 
## 
##  G-structure:  ~us(trait):species
## 
##                                    post.mean  l-95% CI  u-95% CI eff.samp
## traitTFTx:traitTFTx.species        1.3616377  0.784321  2.015956    12667
## traitBPP.CNP:traitTFTx.species    -0.1703082 -0.472095  0.125337    12667
## traitCRISP:traitTFTx.species      -0.0234210 -0.182938  0.119094    12667
## traitCTL:traitTFTx.species        -0.1674153 -0.402719  0.061635    12111
## traitGF:traitTFTx.species          0.0150122 -0.143150  0.177099    12667
## traitKSPI:traitTFTx.species        0.0437850 -0.239427  0.333746    12667
## traitLAAO:traitTFTx.species        0.0120451 -0.116223  0.132538    12667
## traitPLA2:traitTFTx.species       -0.0285271 -0.529122  0.450848    11539
## traitSVMP:traitTFTx.species       -0.6454405 -1.418157  0.016823    12667
## traitSVSP:traitTFTx.species       -0.3970668 -0.850871  0.031453    12667
## traitTFTx:traitBPP.CNP.species    -0.1703082 -0.472095  0.125337    12667
## traitBPP.CNP:traitBPP.CNP.species  0.1910960  0.007720  0.484166    10578
## traitCRISP:traitBPP.CNP.species    0.0169957 -0.089133  0.119452    10086
## traitCTL:traitBPP.CNP.species     -0.0037102 -0.194404  0.175578    10833
## traitGF:traitBPP.CNP.species       0.0126497 -0.087418  0.113557    12667
## traitKSPI:traitBPP.CNP.species    -0.0532846 -0.316200  0.213663    10540
## traitLAAO:traitBPP.CNP.species     0.0003399 -0.080196  0.075905    11187
## traitPLA2:traitBPP.CNP.species     0.0585910 -0.338449  0.500656     8929
## traitSVMP:traitBPP.CNP.species    -0.1574699 -0.836328  0.486297     8292
## traitSVSP:traitBPP.CNP.species     0.1053134 -0.288165  0.468378     9534
## traitTFTx:traitCRISP.species      -0.0234210 -0.182938  0.119094    12667
## traitBPP.CNP:traitCRISP.species    0.0169957 -0.089133  0.119452    10086
## traitCRISP:traitCRISP.species      0.1268462  0.063443  0.198771    12667
## traitCTL:traitCRISP.species       -0.0092549 -0.100461  0.080020    11259
## traitGF:traitCRISP.species         0.0667903  0.009978  0.129071    12667
## traitKSPI:traitCRISP.species      -0.0009373 -0.096469  0.099488    12667
## traitLAAO:traitCRISP.species       0.0661010  0.022287  0.114568    12667
## traitPLA2:traitCRISP.species      -0.1277201 -0.315651  0.030130    11720
## traitSVMP:traitCRISP.species      -0.0537543 -0.284424  0.183855    11890
## traitSVSP:traitCRISP.species      -0.0614364 -0.219417  0.088724     9758
## traitTFTx:traitCTL.species        -0.1674153 -0.402719  0.061635    12111
## traitBPP.CNP:traitCTL.species     -0.0037102 -0.194404  0.175578    10833
## traitCRISP:traitCTL.species       -0.0092549 -0.100461  0.080020    11259
## traitCTL:traitCTL.species          0.1570965  0.011461  0.376445    10911
## traitGF:traitCTL.species           0.0041219 -0.086359  0.087279    12330
## traitKSPI:traitCTL.species         0.1279177 -0.083097  0.342409    10694
## traitLAAO:traitCTL.species         0.0211538 -0.046034  0.082645    11993
## traitPLA2:traitCTL.species        -0.0991896 -0.466568  0.219735    10071
## traitSVMP:traitCTL.species         0.0179580 -0.464173  0.518533     9146
## traitSVSP:traitCTL.species        -0.0485457 -0.369492  0.244654    10125
## traitTFTx:traitGF.species          0.0150122 -0.143150  0.177099    12667
## traitBPP.CNP:traitGF.species       0.0126497 -0.087418  0.113557    12667
## traitCRISP:traitGF.species         0.0667903  0.009978  0.129071    12667
## traitCTL:traitGF.species           0.0041219 -0.086359  0.087279    12330
## traitGF:traitGF.species            0.1084183  0.045592  0.184355    12667
## traitKSPI:traitGF.species          0.0233746 -0.096955  0.141649    12101
## traitLAAO:traitGF.species          0.0673522  0.021137  0.118470    12788
## traitPLA2:traitGF.species         -0.0810327 -0.266094  0.100574    11910
## traitSVMP:traitGF.species         -0.2021413 -0.489541  0.066373    12042
## traitSVSP:traitGF.species         -0.0143324 -0.188095  0.154494    11672
## traitTFTx:traitKSPI.species        0.0437850 -0.239427  0.333746    12667
## traitBPP.CNP:traitKSPI.species    -0.0532846 -0.316200  0.213663    10540
## traitCRISP:traitKSPI.species      -0.0009373 -0.096469  0.099488    12667
## traitCTL:traitKSPI.species         0.1279177 -0.083097  0.342409    10694
## traitGF:traitKSPI.species          0.0233746 -0.096955  0.141649    12101
## traitKSPI:traitKSPI.species        0.4798996  0.243582  0.744500    12667
## traitLAAO:traitKSPI.species        0.0587536 -0.020688  0.145475    12128
## traitPLA2:traitKSPI.species       -0.1905420 -0.575689  0.186106     9989
## traitSVMP:traitKSPI.species       -0.2971399 -0.760485  0.137084    12070
## traitSVSP:traitKSPI.species       -0.1916152 -0.546369  0.131981    10526
## traitTFTx:traitLAAO.species        0.0120451 -0.116223  0.132538    12667
## traitBPP.CNP:traitLAAO.species     0.0003399 -0.080196  0.075905    11187
## traitCRISP:traitLAAO.species       0.0661010  0.022287  0.114568    12667
## traitCTL:traitLAAO.species         0.0211538 -0.046034  0.082645    11993
## traitGF:traitLAAO.species          0.0673522  0.021137  0.118470    12788
## traitKSPI:traitLAAO.species        0.0587536 -0.020688  0.145475    12128
## traitLAAO:traitLAAO.species        0.0834478  0.042730  0.131650    12667
## traitPLA2:traitLAAO.species       -0.1284485 -0.274754 -0.002873    12336
## traitSVMP:traitLAAO.species       -0.0914524 -0.284819  0.107506    12667
## traitSVSP:traitLAAO.species       -0.0891038 -0.222068  0.022852    11957
## traitTFTx:traitPLA2.species       -0.0285271 -0.529122  0.450848    11539
## traitBPP.CNP:traitPLA2.species     0.0585910 -0.338449  0.500656     8929
## traitCRISP:traitPLA2.species      -0.1277201 -0.315651  0.030130    11720
## traitCTL:traitPLA2.species        -0.0991896 -0.466568  0.219735    10071
## traitGF:traitPLA2.species         -0.0810327 -0.266094  0.100574    11910
## traitKSPI:traitPLA2.species       -0.1905420 -0.575689  0.186106     9989
## traitLAAO:traitPLA2.species       -0.1284485 -0.274754 -0.002873    12336
## traitPLA2:traitPLA2.species        0.9011134  0.106816  1.903411     8472
## traitSVMP:traitPLA2.species       -0.8134363 -1.916516  0.206631     7822
## traitSVSP:traitPLA2.species        0.5094027 -0.028889  1.115408     9416
## traitTFTx:traitSVMP.species       -0.6454405 -1.418157  0.016823    12667
## traitBPP.CNP:traitSVMP.species    -0.1574699 -0.836328  0.486297     8292
## traitCRISP:traitSVMP.species      -0.0537543 -0.284424  0.183855    11890
## traitCTL:traitSVMP.species         0.0179580 -0.464173  0.518533     9146
## traitGF:traitSVMP.species         -0.2021413 -0.489541  0.066373    12042
## traitKSPI:traitSVMP.species       -0.2971399 -0.760485  0.137084    12070
## traitLAAO:traitSVMP.species       -0.0914524 -0.284819  0.107506    12667
## traitPLA2:traitSVMP.species       -0.8134363 -1.916516  0.206631     7822
## traitSVMP:traitSVMP.species        2.8078732  1.465037  4.403863    10729
## traitSVSP:traitSVMP.species       -0.5649643 -1.457827  0.232311     8787
## traitTFTx:traitSVSP.species       -0.3970668 -0.850871  0.031453    12667
## traitBPP.CNP:traitSVSP.species     0.1053134 -0.288165  0.468378     9534
## traitCRISP:traitSVSP.species      -0.0614364 -0.219417  0.088724     9758
## traitCTL:traitSVSP.species        -0.0485457 -0.369492  0.244654    10125
## traitGF:traitSVSP.species         -0.0143324 -0.188095  0.154494    11672
## traitKSPI:traitSVSP.species       -0.1916152 -0.546369  0.131981    10526
## traitLAAO:traitSVSP.species       -0.0891038 -0.222068  0.022852    11957
## traitPLA2:traitSVSP.species        0.5094027 -0.028889  1.115408     9416
## traitSVMP:traitSVSP.species       -0.5649643 -1.457827  0.232311     8787
## traitSVSP:traitSVSP.species        0.7525326  0.108657  1.517038     8665
## 
##  R-structure:  ~us(trait):units
## 
##                                  post.mean  l-95% CI  u-95% CI eff.samp
## traitTFTx:traitTFTx.units        0.0858214  0.011802  0.187894    12667
## traitBPP.CNP:traitTFTx.units     0.0460542 -0.120860  0.208354    12212
## traitCRISP:traitTFTx.units      -0.0181114 -0.053798  0.013516    12667
## traitCTL:traitTFTx.units        -0.0143573 -0.131474  0.098669    12667
## traitGF:traitTFTx.units         -0.0203481 -0.073116  0.031104    12064
## traitKSPI:traitTFTx.units       -0.0160058 -0.075472  0.038938    12008
## traitLAAO:traitTFTx.units       -0.0007266 -0.031143  0.027869    12667
## traitPLA2:traitTFTx.units       -0.1601139 -0.373440  0.032368    11680
## traitSVMP:traitTFTx.units        0.0128448 -0.122675  0.139115    11675
## traitSVSP:traitTFTx.units        0.0850575 -0.089988  0.264240    13241
## traitTFTx:traitBPP.CNP.units     0.0460542 -0.120860  0.208354    12212
## traitBPP.CNP:traitBPP.CNP.units  0.5309996  0.296460  0.815570    12595
## traitCRISP:traitBPP.CNP.units   -0.0537672 -0.118242  0.005450    12677
## traitCTL:traitBPP.CNP.units     -0.0241815 -0.156985  0.101900    12667
## traitGF:traitBPP.CNP.units      -0.0488238 -0.123680  0.025294    15193
## traitKSPI:traitBPP.CNP.units    -0.0584180 -0.177029  0.058894    11170
## traitLAAO:traitBPP.CNP.units    -0.0380964 -0.090020  0.012244    12667
## traitPLA2:traitBPP.CNP.units    -0.1553666 -0.446966  0.110078    11019
## traitSVMP:traitBPP.CNP.units    -0.0354619 -0.312299  0.229912     9933
## traitSVSP:traitBPP.CNP.units    -0.1628645 -0.407546  0.075413    12192
## traitTFTx:traitCRISP.units      -0.0181114 -0.053798  0.013516    12667
## traitBPP.CNP:traitCRISP.units   -0.0537672 -0.118242  0.005450    12677
## traitCRISP:traitCRISP.units      0.0220577  0.005732  0.042005    12667
## traitCTL:traitCRISP.units       -0.0174918 -0.061664  0.027076    12004
## traitGF:traitCRISP.units         0.0177173 -0.003259  0.042464    12667
## traitKSPI:traitCRISP.units       0.0090654 -0.015784  0.037398    12667
## traitLAAO:traitCRISP.units       0.0058935 -0.006080  0.020701    12667
## traitPLA2:traitCRISP.units       0.0617153 -0.015758  0.143867    13007
## traitSVMP:traitCRISP.units      -0.0078133 -0.067635  0.046747    11914
## traitSVSP:traitCRISP.units      -0.0191603 -0.086568  0.051515    12138
## traitTFTx:traitCTL.units        -0.0143573 -0.131474  0.098669    12667
## traitBPP.CNP:traitCTL.units     -0.0241815 -0.156985  0.101900    12667
## traitCRISP:traitCTL.units       -0.0174918 -0.061664  0.027076    12004
## traitCTL:traitCTL.units          0.2072723  0.088085  0.344203    11776
## traitGF:traitCTL.units          -0.0215050 -0.074257  0.028829    12346
## traitKSPI:traitCTL.units        -0.0106628 -0.097506  0.082617    10952
## traitLAAO:traitCTL.units         0.0086916 -0.024732  0.044895    12667
## traitPLA2:traitCTL.units        -0.0358730 -0.228342  0.152856    10740
## traitSVMP:traitCTL.units         0.0560945 -0.141272  0.261587     9329
## traitSVSP:traitCTL.units        -0.1478923 -0.333133  0.020358    11415
## traitTFTx:traitGF.units         -0.0203481 -0.073116  0.031104    12064
## traitBPP.CNP:traitGF.units      -0.0488238 -0.123680  0.025294    15193
## traitCRISP:traitGF.units         0.0177173 -0.003259  0.042464    12667
## traitCTL:traitGF.units          -0.0215050 -0.074257  0.028829    12346
## traitGF:traitGF.units            0.0587866  0.023327  0.098969    12667
## traitKSPI:traitGF.units          0.0081789 -0.036202  0.047139    11920
## traitLAAO:traitGF.units         -0.0118979 -0.028628  0.005252    12667
## traitPLA2:traitGF.units          0.0190060 -0.092027  0.117440    12065
## traitSVMP:traitGF.units          0.0028874 -0.082835  0.086527    11193
## traitSVSP:traitGF.units         -0.0039022 -0.090642  0.090608    12667
## traitTFTx:traitKSPI.units       -0.0160058 -0.075472  0.038938    12008
## traitBPP.CNP:traitKSPI.units    -0.0584180 -0.177029  0.058894    11170
## traitCRISP:traitKSPI.units       0.0090654 -0.015784  0.037398    12667
## traitCTL:traitKSPI.units        -0.0106628 -0.097506  0.082617    10952
## traitGF:traitKSPI.units          0.0081789 -0.036202  0.047139    11920
## traitKSPI:traitKSPI.units        0.0475200  0.003544  0.112618    11891
## traitLAAO:traitKSPI.units        0.0067292 -0.016180  0.031338    12667
## traitPLA2:traitKSPI.units        0.0648489 -0.086420  0.212931    11232
## traitSVMP:traitKSPI.units       -0.0272311 -0.140801  0.069558    10457
## traitSVSP:traitKSPI.units       -0.0239376 -0.153571  0.104458    10804
## traitTFTx:traitLAAO.units       -0.0007266 -0.031143  0.027869    12667
## traitBPP.CNP:traitLAAO.units    -0.0380964 -0.090020  0.012244    12667
## traitCRISP:traitLAAO.units       0.0058935 -0.006080  0.020701    12667
## traitCTL:traitLAAO.units         0.0086916 -0.024732  0.044895    12667
## traitGF:traitLAAO.units         -0.0118979 -0.028628  0.005252    12667
## traitKSPI:traitLAAO.units        0.0067292 -0.016180  0.031338    12667
## traitLAAO:traitLAAO.units        0.0170905  0.003755  0.032872    12667
## traitPLA2:traitLAAO.units        0.0236237 -0.039577  0.089603    12115
## traitSVMP:traitLAAO.units       -0.0011884 -0.051198  0.046016    12134
## traitSVSP:traitLAAO.units       -0.0100132 -0.074498  0.044650    12044
## traitTFTx:traitPLA2.units       -0.1601139 -0.373440  0.032368    11680
## traitBPP.CNP:traitPLA2.units    -0.1553666 -0.446966  0.110078    11019
## traitCRISP:traitPLA2.units       0.0617153 -0.015758  0.143867    13007
## traitCTL:traitPLA2.units        -0.0358730 -0.228342  0.152856    10740
## traitGF:traitPLA2.units          0.0190060 -0.092027  0.117440    12065
## traitKSPI:traitPLA2.units        0.0648489 -0.086420  0.212931    11232
## traitLAAO:traitPLA2.units        0.0236237 -0.039577  0.089603    12115
## traitPLA2:traitPLA2.units        0.8643911  0.382110  1.407766    10754
## traitSVMP:traitPLA2.units       -0.2388502 -0.634400  0.103320     8935
## traitSVSP:traitPLA2.units       -0.4432701 -0.805764 -0.107121    11066
## traitTFTx:traitSVMP.units        0.0128448 -0.122675  0.139115    11675
## traitBPP.CNP:traitSVMP.units    -0.0354619 -0.312299  0.229912     9933
## traitCRISP:traitSVMP.units      -0.0078133 -0.067635  0.046747    11914
## traitCTL:traitSVMP.units         0.0560945 -0.141272  0.261587     9329
## traitGF:traitSVMP.units          0.0028874 -0.082835  0.086527    11193
## traitKSPI:traitSVMP.units       -0.0272311 -0.140801  0.069558    10457
## traitLAAO:traitSVMP.units       -0.0011884 -0.051198  0.046016    12134
## traitPLA2:traitSVMP.units       -0.2388502 -0.634400  0.103320     8935
## traitSVMP:traitSVMP.units        0.2099677  0.005995  0.568893     9408
## traitSVSP:traitSVMP.units        0.0288413 -0.278611  0.315450     9386
## traitTFTx:traitSVSP.units        0.0850575 -0.089988  0.264240    13241
## traitBPP.CNP:traitSVSP.units    -0.1628645 -0.407546  0.075413    12192
## traitCRISP:traitSVSP.units      -0.0191603 -0.086568  0.051515    12138
## traitCTL:traitSVSP.units        -0.1478923 -0.333133  0.020358    11415
## traitGF:traitSVSP.units         -0.0039022 -0.090642  0.090608    12667
## traitKSPI:traitSVSP.units       -0.0239376 -0.153571  0.104458    10804
## traitLAAO:traitSVSP.units       -0.0100132 -0.074498  0.044650    12044
## traitPLA2:traitSVSP.units       -0.4432701 -0.805764 -0.107121    11066
## traitSVMP:traitSVSP.units        0.0288413 -0.278611  0.315450     9386
## traitSVSP:traitSVSP.units        0.6972702  0.330560  1.126846    11224
## 
##  Location effects: cbind(TFTx, BPP.CNP, CRISP, CTL, GF, KSPI, LAAO, PLA2, SVMP, SVSP) ~ trait:1 
## 
##              post.mean   l-95% CI   u-95% CI eff.samp pMCMC
## (Intercept) -3.085e-05 -6.091e-03  5.929e-03    12667 0.993
posterior.mode(cov_model.1$VCV)
##       traitTFTx:traitTFTx.species    traitBPP.CNP:traitTFTx.species 
##                      1.296487e+00                     -1.489847e-01 
##      traitCRISP:traitTFTx.species        traitCTL:traitTFTx.species 
##                     -7.984542e-03                     -1.278094e-01 
##         traitGF:traitTFTx.species       traitKSPI:traitTFTx.species 
##                      2.693729e-02                     -9.583644e-03 
##       traitLAAO:traitTFTx.species       traitPLA2:traitTFTx.species 
##                     -7.576171e-05                      2.490650e-02 
##       traitSVMP:traitTFTx.species       traitSVSP:traitTFTx.species 
##                     -5.536074e-01                     -3.441671e-01 
##    traitTFTx:traitBPP.CNP.species traitBPP.CNP:traitBPP.CNP.species 
##                     -1.489847e-01                      6.983069e-02 
##   traitCRISP:traitBPP.CNP.species     traitCTL:traitBPP.CNP.species 
##                      2.522745e-02                      9.793705e-03 
##      traitGF:traitBPP.CNP.species    traitKSPI:traitBPP.CNP.species 
##                      9.667959e-03                     -3.401396e-02 
##    traitLAAO:traitBPP.CNP.species    traitPLA2:traitBPP.CNP.species 
##                      8.783221e-03                      6.538883e-02 
##    traitSVMP:traitBPP.CNP.species    traitSVSP:traitBPP.CNP.species 
##                     -6.350128e-02                      1.138660e-01 
##      traitTFTx:traitCRISP.species   traitBPP.CNP:traitCRISP.species 
##                     -7.984542e-03                      2.522745e-02 
##     traitCRISP:traitCRISP.species       traitCTL:traitCRISP.species 
##                      1.048861e-01                      4.493577e-04 
##        traitGF:traitCRISP.species      traitKSPI:traitCRISP.species 
##                      5.746307e-02                     -1.078678e-02 
##      traitLAAO:traitCRISP.species      traitPLA2:traitCRISP.species 
##                      5.684165e-02                     -1.097994e-01 
##      traitSVMP:traitCRISP.species      traitSVSP:traitCRISP.species 
##                     -6.450149e-02                     -4.897689e-02 
##        traitTFTx:traitCTL.species     traitBPP.CNP:traitCTL.species 
##                     -1.278094e-01                      9.793705e-03 
##       traitCRISP:traitCTL.species         traitCTL:traitCTL.species 
##                      4.493577e-04                      8.726539e-02 
##          traitGF:traitCTL.species        traitKSPI:traitCTL.species 
##                      1.233212e-02                      9.986601e-02 
##        traitLAAO:traitCTL.species        traitPLA2:traitCTL.species 
##                      1.417658e-02                     -5.742939e-02 
##        traitSVMP:traitCTL.species        traitSVSP:traitCTL.species 
##                      1.008568e-02                      1.426480e-02 
##         traitTFTx:traitGF.species      traitBPP.CNP:traitGF.species 
##                      2.693729e-02                      9.667959e-03 
##        traitCRISP:traitGF.species          traitCTL:traitGF.species 
##                      5.746307e-02                      1.233212e-02 
##           traitGF:traitGF.species         traitKSPI:traitGF.species 
##                      8.785163e-02                      2.742054e-02 
##         traitLAAO:traitGF.species         traitPLA2:traitGF.species 
##                      5.996534e-02                     -5.742063e-02 
##         traitSVMP:traitGF.species         traitSVSP:traitGF.species 
##                     -2.092450e-01                     -1.924070e-02 
##       traitTFTx:traitKSPI.species    traitBPP.CNP:traitKSPI.species 
##                     -9.583644e-03                     -3.401396e-02 
##      traitCRISP:traitKSPI.species        traitCTL:traitKSPI.species 
##                     -1.078678e-02                      9.986601e-02 
##         traitGF:traitKSPI.species       traitKSPI:traitKSPI.species 
##                      2.742054e-02                      4.652637e-01 
##       traitLAAO:traitKSPI.species       traitPLA2:traitKSPI.species 
##                      5.509807e-02                     -1.855221e-01 
##       traitSVMP:traitKSPI.species       traitSVSP:traitKSPI.species 
##                     -2.870425e-01                     -1.646642e-01 
##       traitTFTx:traitLAAO.species    traitBPP.CNP:traitLAAO.species 
##                     -7.576171e-05                      8.783221e-03 
##      traitCRISP:traitLAAO.species        traitCTL:traitLAAO.species 
##                      5.684165e-02                      1.417658e-02 
##         traitGF:traitLAAO.species       traitKSPI:traitLAAO.species 
##                      5.996534e-02                      5.509807e-02 
##       traitLAAO:traitLAAO.species       traitPLA2:traitLAAO.species 
##                      7.579598e-02                     -1.091065e-01 
##       traitSVMP:traitLAAO.species       traitSVSP:traitLAAO.species 
##                     -7.223834e-02                     -6.580230e-02 
##       traitTFTx:traitPLA2.species    traitBPP.CNP:traitPLA2.species 
##                      2.490650e-02                      6.538883e-02 
##      traitCRISP:traitPLA2.species        traitCTL:traitPLA2.species 
##                     -1.097994e-01                     -5.742939e-02 
##         traitGF:traitPLA2.species       traitKSPI:traitPLA2.species 
##                     -5.742063e-02                     -1.855221e-01 
##       traitLAAO:traitPLA2.species       traitPLA2:traitPLA2.species 
##                     -1.091065e-01                      7.216318e-01 
##       traitSVMP:traitPLA2.species       traitSVSP:traitPLA2.species 
##                     -6.697253e-01                      4.814302e-01 
##       traitTFTx:traitSVMP.species    traitBPP.CNP:traitSVMP.species 
##                     -5.536074e-01                     -6.350128e-02 
##      traitCRISP:traitSVMP.species        traitCTL:traitSVMP.species 
##                     -6.450149e-02                      1.008568e-02 
##         traitGF:traitSVMP.species       traitKSPI:traitSVMP.species 
##                     -2.092450e-01                     -2.870425e-01 
##       traitLAAO:traitSVMP.species       traitPLA2:traitSVMP.species 
##                     -7.223834e-02                     -6.697253e-01 
##       traitSVMP:traitSVMP.species       traitSVSP:traitSVMP.species 
##                      2.644246e+00                     -4.618826e-01 
##       traitTFTx:traitSVSP.species    traitBPP.CNP:traitSVSP.species 
##                     -3.441671e-01                      1.138660e-01 
##      traitCRISP:traitSVSP.species        traitCTL:traitSVSP.species 
##                     -4.897689e-02                      1.426480e-02 
##         traitGF:traitSVSP.species       traitKSPI:traitSVSP.species 
##                     -1.924070e-02                     -1.646642e-01 
##       traitLAAO:traitSVSP.species       traitPLA2:traitSVSP.species 
##                     -6.580230e-02                      4.814302e-01 
##       traitSVMP:traitSVSP.species       traitSVSP:traitSVSP.species 
##                     -4.618826e-01                      5.643247e-01 
##         traitTFTx:traitTFTx.units      traitBPP.CNP:traitTFTx.units 
##                      5.531391e-02                      2.015981e-02 
##        traitCRISP:traitTFTx.units          traitCTL:traitTFTx.units 
##                     -1.661541e-02                     -1.075508e-03 
##           traitGF:traitTFTx.units         traitKSPI:traitTFTx.units 
##                     -1.133672e-02                     -1.489698e-02 
##         traitLAAO:traitTFTx.units         traitPLA2:traitTFTx.units 
##                     -5.644959e-04                     -1.226980e-01 
##         traitSVMP:traitTFTx.units         traitSVSP:traitTFTx.units 
##                      1.006565e-02                      9.709177e-02 
##      traitTFTx:traitBPP.CNP.units   traitBPP.CNP:traitBPP.CNP.units 
##                      2.015981e-02                      4.863293e-01 
##     traitCRISP:traitBPP.CNP.units       traitCTL:traitBPP.CNP.units 
##                     -5.090489e-02                     -2.932575e-02 
##        traitGF:traitBPP.CNP.units      traitKSPI:traitBPP.CNP.units 
##                     -4.814097e-02                     -5.457160e-02 
##      traitLAAO:traitBPP.CNP.units      traitPLA2:traitBPP.CNP.units 
##                     -4.439142e-02                     -1.032140e-01 
##      traitSVMP:traitBPP.CNP.units      traitSVSP:traitBPP.CNP.units 
##                     -8.419889e-03                     -1.312664e-01 
##        traitTFTx:traitCRISP.units     traitBPP.CNP:traitCRISP.units 
##                     -1.661541e-02                     -5.090489e-02 
##       traitCRISP:traitCRISP.units         traitCTL:traitCRISP.units 
##                      1.765437e-02                     -1.650449e-02 
##          traitGF:traitCRISP.units        traitKSPI:traitCRISP.units 
##                      1.217460e-02                      9.489824e-03 
##        traitLAAO:traitCRISP.units        traitPLA2:traitCRISP.units 
##                      1.981300e-03                      5.060888e-02 
##        traitSVMP:traitCRISP.units        traitSVSP:traitCRISP.units 
##                     -1.895829e-03                     -1.232197e-02 
##          traitTFTx:traitCTL.units       traitBPP.CNP:traitCTL.units 
##                     -1.075508e-03                     -2.932575e-02 
##         traitCRISP:traitCTL.units           traitCTL:traitCTL.units 
##                     -1.650449e-02                      1.847474e-01 
##            traitGF:traitCTL.units          traitKSPI:traitCTL.units 
##                     -2.265472e-02                     -2.184210e-02 
##          traitLAAO:traitCTL.units          traitPLA2:traitCTL.units 
##                      8.427578e-03                     -2.396657e-02 
##          traitSVMP:traitCTL.units          traitSVSP:traitCTL.units 
##                      3.756037e-02                     -1.172093e-01 
##           traitTFTx:traitGF.units        traitBPP.CNP:traitGF.units 
##                     -1.133672e-02                     -4.814097e-02 
##          traitCRISP:traitGF.units            traitCTL:traitGF.units 
##                      1.217460e-02                     -2.265472e-02 
##             traitGF:traitGF.units           traitKSPI:traitGF.units 
##                      4.754728e-02                      1.149869e-02 
##           traitLAAO:traitGF.units           traitPLA2:traitGF.units 
##                     -1.185429e-02                      1.907763e-02 
##           traitSVMP:traitGF.units           traitSVSP:traitGF.units 
##                      3.100707e-03                     -3.504329e-04 
##         traitTFTx:traitKSPI.units      traitBPP.CNP:traitKSPI.units 
##                     -1.489698e-02                     -5.457160e-02 
##        traitCRISP:traitKSPI.units          traitCTL:traitKSPI.units 
##                      9.489824e-03                     -2.184210e-02 
##           traitGF:traitKSPI.units         traitKSPI:traitKSPI.units 
##                      1.149869e-02                      2.786758e-02 
##         traitLAAO:traitKSPI.units         traitPLA2:traitKSPI.units 
##                      5.272555e-03                      7.708143e-02 
##         traitSVMP:traitKSPI.units         traitSVSP:traitKSPI.units 
##                     -9.721501e-03                     -3.383246e-02 
##         traitTFTx:traitLAAO.units      traitBPP.CNP:traitLAAO.units 
##                     -5.644959e-04                     -4.439142e-02 
##        traitCRISP:traitLAAO.units          traitCTL:traitLAAO.units 
##                      1.981300e-03                      8.427578e-03 
##           traitGF:traitLAAO.units         traitKSPI:traitLAAO.units 
##                     -1.185429e-02                      5.272555e-03 
##         traitLAAO:traitLAAO.units         traitPLA2:traitLAAO.units 
##                      1.446954e-02                      2.765632e-02 
##         traitSVMP:traitLAAO.units         traitSVSP:traitLAAO.units 
##                     -3.932400e-04                     -2.399112e-03 
##         traitTFTx:traitPLA2.units      traitBPP.CNP:traitPLA2.units 
##                     -1.226980e-01                     -1.032140e-01 
##        traitCRISP:traitPLA2.units          traitCTL:traitPLA2.units 
##                      5.060888e-02                     -2.396657e-02 
##           traitGF:traitPLA2.units         traitKSPI:traitPLA2.units 
##                      1.907763e-02                      7.708143e-02 
##         traitLAAO:traitPLA2.units         traitPLA2:traitPLA2.units 
##                      2.765632e-02                      7.429985e-01 
##         traitSVMP:traitPLA2.units         traitSVSP:traitPLA2.units 
##                     -1.653414e-01                     -4.225168e-01 
##         traitTFTx:traitSVMP.units      traitBPP.CNP:traitSVMP.units 
##                      1.006565e-02                     -8.419889e-03 
##        traitCRISP:traitSVMP.units          traitCTL:traitSVMP.units 
##                     -1.895829e-03                      3.756037e-02 
##           traitGF:traitSVMP.units         traitKSPI:traitSVMP.units 
##                      3.100707e-03                     -9.721501e-03 
##         traitLAAO:traitSVMP.units         traitPLA2:traitSVMP.units 
##                     -3.932400e-04                     -1.653414e-01 
##         traitSVMP:traitSVMP.units         traitSVSP:traitSVMP.units 
##                      4.850233e-02                      4.620712e-02 
##         traitTFTx:traitSVSP.units      traitBPP.CNP:traitSVSP.units 
##                      9.709177e-02                     -1.312664e-01 
##        traitCRISP:traitSVSP.units          traitCTL:traitSVSP.units 
##                     -1.232197e-02                     -1.172093e-01 
##           traitGF:traitSVSP.units         traitKSPI:traitSVSP.units 
##                     -3.504329e-04                     -3.383246e-02 
##         traitLAAO:traitSVSP.units         traitPLA2:traitSVSP.units 
##                     -2.399112e-03                     -4.225168e-01 
##         traitSVMP:traitSVSP.units         traitSVSP:traitSVSP.units 
##                      4.620712e-02                      5.856831e-01

Supplementary Figure 1: Convergence of MCMC for fixed & random effect

plot(cov_model.1$Sol)