Belowground regulation of
aboveground dynamics
Gaurav Kandlikar
Louisiana State University
web: https://gklab.org/
slides: https://talks.gklab.org/utep-26
contact: gkandlikar@lsu.edu
Sal (Shorea) dominated forests in northern India, Photo by Dhritiman Mukherjee
Intercropping of soybean and wheat; image from Zalf
“Christmas tree scientists work to manage grinchy fungal foes”; image from Washington State Univ
As an ecologist, my research is motivated by two overarching goals:
Explain patterns of diversity in nature
Predict responses to perturbations



Advancing ecological theory and experiments to quantify microbial effects on plant coexistence
Microbial effects on plant communities in variable environments: Ongoing work and new directions
Mathematical underpinning of plant–soil feedback
See also Bever, Westover, and Antonovics (1997) and Kandlikar (2024)
\[\frac{dN_1}{dt} = w_1N_1 ~~~~ \text{and} ~~~ \frac{dN_2}{dt} = w_2N_2\]
\[w_i = m_{iA}f_A + m_{iB}(1-f_A)\]
where \(f_i\) is the relative frequency of microbial community \(i\) in the soil (e.g. \(f_A = \frac{N_A}{N_A+N_B}\))
Interpretation: Plants have exponential population growth at a rate determined by the composition of the soil community.
\[\frac{dN_A}{dt} = N_A\frac{N_1}{N_1 + N_2}~~~\text{and}~~~\frac{dN_B}{dt} = \nu N_B\frac{N_2}{N_1 + N_2}\]
Interpretation: Soil communities grow at a rate determined by the frequency of each plant (and the relative strength of each plant’s conditioning effect, \(\nu\)).
R code to simulate model dynamics
psf_model <- function(time, init, params) {
with (as.list(c(time, init, params)), {
# description of parameters (see Bever et al. 1997)
# N1 and N2: abundance of the plant species 1 and 2
# p1: frequency of plant species 1; p2 = 1-pA
# m1A, m1B, m2A, m2B: conspecific and heterospecific effects of microbial community A or B on the growth of plant 1 or 2
# pA: frequency of the soil microbial community A
# v: influence of plant species 2 on the microbial community relative to that of plant 1
# Differential equations
dN1 <- (m1A*pA + m1B*(1-pA))*N1
dN2 <- (m2A*pA + m2B*(1-pA))*N2
dp1 <- p1*(1-p1)*((m1A-m2A)*pA + (m1B-m2B)*(1-pA))
dpA <- pA*(1-pA)*(p1-v*(1-p1))
# Return dN1 and dN2
return(list(c(dN1, dN2, dp1, dpA)))
})
}\[ \overbrace{(m_{1B} + m_{2A})}^{\text{interspecific effects}}- \overbrace{(m_{1A} + m_{2B})}^{\text{intraspecific effects}} > 0 \]
\[ \underbrace{\frac{1}{2}\big[\overbrace{(m_{1B} + m_{2A})}^{\text{interspecific effects}}- \overbrace{(m_{1A} + m_{2B})}^{\text{intraspecific effects}}\big]}_{\text{Stablization}} > 0 \]
# Dataset downloaded from Figshare:
# curl::curl_download("https://figshare.com/ndownloader/files/14874749", destfile = "../crawford-supplement.xlsx")
read_excel("../data/crawford-supplement.xlsx", sheet = "Data") |>
mutate(stab = -0.5*rrIs) |> # show in terms of stabilization instead of I_s
filter(stab > -3) |> # Remove the pair with Stab approx. 6 -- according to McCarthy Neuman (original author), this is likely due to bad seed survival
ggplot(aes(x = stab)) +
geom_histogram(color = "black") +
geom_histogram(fill = "#56A0D3") +
geom_rug(color = "grey25") +
geom_vline(xintercept = 0) +
annotate("text", x = Inf, y = Inf, label = "N = 1037 pairwise comparisons\n
57% experience positive stabilization",
vjust=1, hjust = 1, size = 6) +
xlab("Stabilization") + ylab("Count")
Data from Crawford et al. (2019)
mangan <-
tribble(~IS, ~abun, ~where,
-0.15504664970313828, 0.7732558139534884, "BCI",
-0.0887192536047498, 1.005813953488372, "BCI",
-0.08583545377438509, 1.4825581395348837, "BCI",
-0.09669211195928756, 1.7267441860465116, "BCI",
-0.04512298558100089, 2.1104651162790695, "BCI",
-0.006276505513146763, 2.4186046511627906, "BCI",
-0.18231106613816006, 0.3003106037506997, "Gigante",
-0.1717428991693779, 1.0202629577384568, "Gigante",
-0.13691306110550813, 1.083509520825865, "Gigante",
-0.1250429718781308, 1.425445047930817, "Gigante",
-0.09753373250550346, 1.9673983333947622, "Gigante") |>
mutate(stab = IS*-0.5)
mangan_rug_blank <-
mangan |>
ggplot(aes(x = stab,y = abun, color = where)) +
# geom_rug(sides = 'b', linewidth = 2) +
# geom_point(size = 4, stroke = 2, shape = 21) +
# geom_smooth(method = "lm", se = F) +
geom_point(color = 'transparent') +
scale_color_manual(values = c("#117733", "#cc6677")) +
geom_vline(xintercept = 0, linetype = 'dashed', color = 'grey', linewidth = 2) +
xlab("Strength of feedback (species' average stablization)") +
ylab("Log abundance of trees") +
theme(legend.position = 'none') +
scale_x_continuous(limits = c(-0.1, 0.1))+
scale_y_continuous(limits = c(-0.1,2.5))
mangan_rug_blank
Data from Mangan et al. (2010)

Data from Mangan et al. (2010)

Data from Mangan et al. (2010)

Data from Mangan et al. (2010)
mangan_rug +
geom_point(size = 4, stroke = 2, shape = 21) +
geom_smooth(method = "lm", se = F) +
annotate('text', x = -0.1, y = 2, fontface = "italic", hjust = 0, vjust = 0.5, size = 5,
label = '"[...] trees are abundant because they are less\nsusceptible to the detrimental effects of\ntheir associated soil communities than are\nrarer tree species"')
Data from Mangan et al. (2010)
# Define a function used for making the PSF framework
# schematic, given a set of parameter values
make_params_plot <- function(params, scale = 1.5) {
color_func <- function(x) {
ifelse(x < 0, "#EE6677", "#4477AA")
}
df <- data.frame(x = c(0,0,1,1),
y = c(0,1,0,1),
type = c("M", "P", "M", "P"))
params_plot <-
ggplot(df) +
annotate("text", x = 0, y = 1.1, size = 3.25, label = "Plant 1",
color = "black", fill = "#9970ab", fontface = "bold") +
annotate("text", x = 1, y = 1.1, size = 3.25, label = "Plant 2",
color = "black", fill = "#5aae61", fontface = "bold") +
annotate("text", x = 0, y = -0.15, size = 3.25,
label = "Soil\nmicrobes A", label.size = 0, fill = "transparent") +
annotate("text", x = 1, y = -0.15, size = 3.25,
label = "Soil\nmicrobes B", label.size = 0, fill = "transparent") +
annotate("text", x = 0.45, y = -0.4, size = 3.5, fill = "transparent",
label.size = 0.05,
label = (paste0("Stabilization = ",
0.5*(params["m1B"] + params["m2A"] - params["m1A"] - params["m2B"])))) +
geom_segment(aes(x = 0, xend = 0, y = 0.1, yend = 0.9),
arrow = arrow(length = unit(0.03, "npc")),
linewidth = abs(params["m1A"])*scale,
color = alpha(color_func(params["m1A"]), 1)) +
geom_segment(aes(x = 0.05, xend = 0.95, y = 0.1, yend = 0.9),
arrow = arrow(length = unit(0.03, "npc")),
linewidth = abs(params["m2A"])*scale,
color = alpha(color_func(params["m1B"]),1)) +
geom_segment(aes(x = 0.95, xend = 0.05, y = 0.1, yend = 0.9),
arrow = arrow(length = unit(0.03, "npc")),
linewidth = abs(params["m1B"])*scale,
color = alpha(color_func(params["m2A"]), 1)) +
geom_segment(aes(x = 1, xend = 1, y = 0.1, yend = 0.9),
arrow = arrow(length = unit(0.03, "npc"),),
linewidth = abs(params["m2B"])*scale,
color = alpha(color_func(params["m2B"]), 1)) +
# Plant cultivation of microbes
geom_segment(aes(x = -0.25, xend = -0.25, y = 0.9, yend = 0.1),
linewidth = 0.15, linetype = 1,
arrow = arrow(length = unit(0.03, "npc"))) +
geom_segment(aes(x = 1.25, xend = 1.25, y = 0.9, yend = 0.1),
linewidth = 0.15, linetype = 1,
arrow = arrow(length = unit(0.03, "npc"))) +
annotate("text", x = 0, y = 0.5,
label = (paste0("m1A = ", params["m1A"])),
angle = 90, vjust = -0.25, size = 3) +
annotate("text", x = 1.15, y = 0.5,
label = (paste0("m2B = ", params["m2B"])),
angle = -90, vjust = 1.5, size = 3) +
annotate("text", x = 0.75, y = 0.75,
label = (paste0("m2A = ", params["m2A"])),
angle = 45, vjust = -0.25, size = 3) +
annotate("text", x = 0.25, y = 0.75,
label = (paste0("m1B = ", params["m1B"])),
angle = -45, vjust = -0.25, size = 3) +
xlim(c(-0.4, 1.4)) +
coord_cartesian(ylim = c(-0.25, 1.15), clip = "off") +
theme_void() +
theme(legend.position = "none",
plot.caption = element_text(hjust = 0.5, size = 10))
return(params_plot)
}
# Define a function for simulating the dynamics of the
# PSF model with deSolve
params_coex <- c(m10 = 0.16, m20 = 0.16,
m1A = 0.11, m1B = 0.26,
m2A = 0.27, m2B = 0.13, v = 1)
time <- seq(0,50,0.1)
init_pA_05 <- c(N1 = 3, N2 = 7, p1 = 0.3, pA = 0.3)
out_pA_05 <- ode(y = init_pA_05, times = time, func = psf_model, parms = params_coex) |> data.frame()
params_to_plot_coex <- c(m1A = unname(params_coex["m1A"]-params_coex["m10"]),
m1B = unname(params_coex["m1B"]-params_coex["m10"]),
m2A = unname(params_coex["m2A"]-params_coex["m20"]),
m2B = unname(params_coex["m2B"]-params_coex["m20"]), v=1)
param_plot_coex <-
make_params_plot(params_to_plot_coex, scale = 6) +
annotate("text", x = 1.375, y = 0.5,
label = paste0("v = ", params_coex["v"]),
angle = -90, vjust = 1.5, size = 3)
panel_abund_coex <-
out_pA_05 |>
as_tibble() |>
select(time, N1, N2) |>
pivot_longer(N1:N2) |>
ggplot(aes(x = time, y = value, color = name)) +
geom_line(linewidth = 0.9) +
scale_color_manual(values = c("#9970ab", "#5aae61"),
name = "Plant species", label = c("Plant 1", "Plant 2"), guide = "none") +
scale_y_continuous(breaks = c(2e4, 4e4, 6e4, 8e4), labels = scales::scientific) +
ylab("Abundance") +
theme(axis.title = element_text(size = 10))
# Panel D: plot for frequencies of both plants when growing
# in dynamic soils
panel_freq_coex <-
out_pA_05 |>
as_tibble() |>
mutate(p2 = 1-p1) |>
select(time, p1, p2) |>
pivot_longer(p1:p2) |>
ggplot(aes(x = time, y = value, color = name)) +
geom_line(linewidth = 1.2) +
scale_color_manual(values = c("#9970ab", "#5aae61"),
name = "Plant species", label = c("Plant 1", "Plant 2")) +
scale_y_continuous(limits = c(0,1), breaks = c(0, 0.5, 1)) +
ylab("Frequency") +
theme(axis.title = element_text(size = 10))
param_plot_coex + {panel_abund_coex+panel_freq_coex &
theme(axis.text = element_text(size = 8), legend.position = 'none')} +
plot_layout(widths = c(1/3, 2/3))
# Define a function for simulating the dynamics of the
# PSF model with deSolve
params <- c(m10 = 0.16, m20 = 0.16,
m1A = 0.02, m2A = 0.33,
m1B = 0.18, m2B = 0.20, v = 1)
time <- seq(0,200,0.1)
init_pA_05 <- c(N1 = 3, N2 = 7, p1 = 0.3, pA = 0.3)
out_pA_05 <- ode(y = init_pA_05, times = time, func = psf_model, parms = params) |> data.frame()
params_to_plot <- c(m1A = unname(params["m1A"]-params["m10"]),
m1B = unname(params["m1B"]-params["m10"]),
m2A = unname(params["m2A"]-params["m20"]),
m2B = unname(params["m2B"]-params["m20"]), v=1)
param_plot <-
make_params_plot(params_to_plot, scale = 6) +
annotate("text", x = 1.375, y = 0.5,
label = paste0("v = ", params["v"]),
angle = -90, vjust = 1.5, size = 3)
panel_abund <-
out_pA_05 |>
as_tibble() |>
select(time, N1, N2) |>
pivot_longer(N1:N2) |>
ggplot(aes(x = time, y = value, color = name)) +
geom_line(linewidth = 0.9) +
scale_color_manual(values = c("#9970ab", "#5aae61"),
name = "Plant species", label = c("Plant 1", "Plant 2"), guide = "none") +
scale_y_continuous(breaks = c(0,8e17,1.6e18), labels = c("0", "7e17","1.4e18")) +
ylab("Abundance")
# Panel D: plot for frequencies of both plants when growing
# in dynamic soils
panel_freq <-
out_pA_05 |>
as_tibble() |>
mutate(p2 = 1-p1) |>
select(time, p1, p2) |>
pivot_longer(p1:p2) |>
ggplot(aes(x = time, y = value, color = name)) +
geom_line(linewidth = 1.2) +
scale_color_manual(values = c("#9970ab", "#5aae61"),
name = "Plant species", label = c("Plant 1", "Plant 2")) +
scale_y_continuous(limits = c(0,1), breaks = c(0, 0.5, 1)) +
ylab("Frequency")
param_plot + {panel_abund + panel_freq &
theme(axis.text = element_text(size = 8), legend.position = 'none')} +
plot_layout(widths = c(1/3, 2/3))
Build on insights from coexistence theory (Chesson 2018):
Stabilization is necessary for coexistence but doesn’t guarantee coexistence
Stable coexistence is possible only when stabilization overcomes competitive imbalances (fitness differences)





\[\text{Fitness difference}_{1,2} = \overbrace{\frac{1}{2}(m_{1A}+m_{1B})}^{\substack{\text{Sensitivity of}\\ \text{Sp 1 to microbes}}} - \overbrace{\frac{1}{2}(m_{2A}+m_{2B})}^{\substack{\text{Sensitivity of}\\ \text{Sp 2 to microbes}}}\]
base <-
tibble(sd = seq(-1,1,0.1),
fd = c(seq(1,0,-0.1), seq(0.1,1,0.1))) |>
ggplot(aes(x = sd, y = fd)) +
geom_line(linetype = "dashed") +
geom_hline(yintercept = 0, linewidth = 1) +
geom_vline(xintercept = 0, linewidth = 1) +
xlab("Stabilization") +
ylab("Fitness difference") +
theme(axis.text = element_blank(),
axis.ticks = element_blank(),
axis.line = element_blank(),
axis.title = element_text(size = 25)) +
scale_x_continuous(expand = c(0,0)) +
scale_y_continuous(expand = c(0.04,0))
base
base <-
base +
annotate("text", x = Inf, y = 0, hjust = 1, vjust = -1,
label = "Coexistence", size = 7, family = "italic") +
annotate("text", x = -Inf, y = 0, hjust = 0, vjust = -1,
label = "Priority effects", size = 7, family = "italic") +
annotate("label", x = 0, y = Inf, vjust = 1.5,
label = "Species exclusion", size = 7, fill = "#F0F1EB", family = "italic")
base
metrics <- function(params) {
IS <- with(as.list(params), {m1A - m2A - m1B + m2B})
FD <- with(as.list(params), {(1/2)*(m1A+m2A) - (1/2)*(m1B+m2B)})
SD <- (-1/2)*IS
return(c(IS = IS, SD = SD, FD = FD))
}
coex_metrics <- metrics(params_coex)
base_coex <-
base +
inset_element({panel_freq_coex + theme(legend.position = 'none',
axis.text = element_blank(),
axis.ticks = element_blank(),
axis.title = element_blank(),
plot.background = element_rect(color = "grey"))},
0.75,0.15,1,0.5)
base_coex
Sedgwick Reserve (Santa Barbara County, California)

# Dataset downloaded from Figshare:
# curl::curl_download("https://datadryad.org/downloads/file_stream/367188", destfile = "../data/kandlikar2021-supplement.csv")
biomass_wide <-
read_csv("../data/kandlikar2021-supplement.csv") |>
mutate(log_agb = log(abg_dry_g)) |>
mutate(source_soil = ifelse(source_soil == "aastr", "str", source_soil),
source_soil = ifelse(source_soil == "abfld", "fld", source_soil),
pair = paste0(source_soil, "_", focal_species)) |>
select(replicate, pair, log_agb) |>
pivot_wider(names_from = pair, values_from = log_agb) |>
filter(!is.na(replicate))
# Calculate the Stabilization between each species pair ----
# Recall that following the definition of the m terms in Bever 1997,
# and the analysis of this model in Kandlikar 2019,
# stablization = -0.5*(log(m1A) + log(m2B) - log(m1B) - log(m2A))
# Here is a function that does this calculation
# (recall that after the data reshaping above,
# each column represents a given value of log(m1A))
calculate_stabilization <- function(df) {
df |>
# In df, each row is one repilcate/rack, and each column
# represents the growth of one species in one soil type.
# e.g. "FE_ACWR" is the growth of FE in ACWR-cultivated soil.
mutate(AC_FE = -0.5*(AC_ACWR - AC_FEMI - FE_ACWR + FE_FEMI),
AC_HO = -0.5*(AC_ACWR - AC_HOMU - HO_ACWR + HO_HOMU),
AC_SA = -0.5*(AC_ACWR - AC_SACO - SA_ACWR + SA_SACO),
AC_PL = -0.5*(AC_ACWR - AC_PLER - PL_ACWR + PL_PLER),
AC_UR = -0.5*(AC_ACWR - AC_URLI - UR_ACWR + UR_URLI),
FE_HO = -0.5*(FE_FEMI - FE_HOMU - HO_FEMI + HO_HOMU),
FE_SA = -0.5*(FE_FEMI - FE_SACO - SA_FEMI + SA_SACO),
FE_PL = -0.5*(FE_FEMI - FE_PLER - PL_FEMI + PL_PLER),
FE_UR = -0.5*(FE_FEMI - FE_URLI - UR_FEMI + UR_URLI),
HO_PL = -0.5*(HO_HOMU - HO_PLER - PL_HOMU + PL_PLER),
HO_SA = -0.5*(HO_HOMU - HO_SACO - SA_HOMU + SA_SACO),
HO_UR = -0.5*(HO_HOMU - HO_URLI - UR_HOMU + UR_URLI),
SA_PL = -0.5*(SA_SACO - SA_PLER - PL_SACO + PL_PLER),
SA_UR = -0.5*(SA_SACO - SA_URLI - UR_SACO + UR_URLI),
PL_UR = -0.5*(PL_PLER - PL_URLI - UR_PLER + UR_URLI)) |>
select(replicate, AC_FE:PL_UR) |>
gather(pair, stabilization, AC_FE:PL_UR)
}
stabilization_values <- calculate_stabilization(biomass_wide)
# Seven values are NA; we can omit these
stabilization_values <- stabilization_values |>
filter(!(is.na(stabilization)))
# Calculating the Fitness difference between each species pair ----
# Similarly, we can now calculate the fitness difference between
# each pair. Recall that FD = 0.5*(log(m1A)+log(m1B)-log(m2A)-log(m2B))
# But recall that here, IT IS IMPORTANT THAT
# m1A = (m1_soilA - m1_fieldSoil)!
# The following function does this calculation:
calculate_fitdiffs <- function(df) {
df |>
mutate(AC_FE = 0.5*((AC_ACWR-fld_ACWR) + (FE_ACWR-fld_ACWR) - (AC_FEMI-fld_FEMI) - (FE_FEMI-fld_FEMI)),
AC_HO = 0.5*((AC_ACWR-fld_ACWR) + (HO_ACWR-fld_ACWR) - (AC_HOMU-fld_HOMU) - (HO_HOMU-fld_HOMU)),
AC_SA = 0.5*((AC_ACWR-fld_ACWR) + (SA_ACWR-fld_ACWR) - (AC_SACO-fld_SACO) - (SA_SACO-fld_SACO)),
AC_PL = 0.5*((AC_ACWR-fld_ACWR) + (PL_ACWR-fld_ACWR) - (AC_PLER-fld_PLER) - (PL_PLER-fld_PLER)),
AC_UR = 0.5*((AC_ACWR-fld_ACWR) + (UR_ACWR-fld_ACWR) - (AC_URLI-fld_URLI) - (UR_URLI-fld_URLI)),
FE_HO = 0.5*((FE_FEMI-fld_FEMI) + (HO_FEMI-fld_FEMI) - (FE_HOMU-fld_HOMU) - (HO_HOMU-fld_HOMU)),
FE_SA = 0.5*((FE_FEMI-fld_FEMI) + (SA_FEMI-fld_FEMI) - (FE_SACO-fld_SACO) - (SA_SACO-fld_SACO)),
FE_PL = 0.5*((FE_FEMI-fld_FEMI) + (PL_FEMI-fld_FEMI) - (FE_PLER-fld_PLER) - (PL_PLER-fld_PLER)),
FE_UR = 0.5*((FE_FEMI-fld_FEMI) + (UR_FEMI-fld_FEMI) - (FE_URLI-fld_URLI) - (UR_URLI-fld_URLI)),
HO_PL = 0.5*((HO_HOMU-fld_HOMU) + (PL_HOMU-fld_HOMU) - (HO_PLER-fld_PLER) - (PL_PLER-fld_PLER)),
HO_SA = 0.5*((HO_HOMU-fld_HOMU) + (SA_HOMU-fld_HOMU) - (HO_SACO-fld_SACO) - (SA_SACO-fld_SACO)),
HO_UR = 0.5*((HO_HOMU-fld_HOMU) + (UR_HOMU-fld_HOMU) - (HO_URLI-fld_URLI) - (UR_URLI-fld_URLI)),
SA_PL = 0.5*((SA_SACO-fld_SACO) + (PL_SACO-fld_SACO) - (SA_PLER-fld_PLER) - (PL_PLER-fld_PLER)),
SA_UR = 0.5*((SA_SACO-fld_SACO) + (UR_SACO-fld_SACO) - (SA_URLI-fld_URLI) - (UR_URLI-fld_URLI)),
PL_UR = 0.5*((PL_PLER-fld_PLER) + (UR_PLER-fld_PLER) - (PL_URLI-fld_URLI) - (UR_URLI-fld_URLI))) |>
select(replicate, AC_FE:PL_UR) |>
gather(pair, fitdiff_fld, AC_FE:PL_UR)
}
fd_values <- calculate_fitdiffs(biomass_wide)
# Twenty-two values are NA; let's omit these.
fd_values <- fd_values |> filter(!(is.na(fitdiff_fld)))
# Now, generate statistical summaries of SD and FD
stabiliation_summary <- stabilization_values |> group_by(pair) |>
summarize(mean_sd = mean(stabilization),
sem_sd = sd(stabilization)/sqrt(n()),
n_sd = n())
fitdiff_summary <- fd_values |> group_by(pair) |>
summarize(mean_fd = mean(fitdiff_fld),
sem_fd = sd(fitdiff_fld)/sqrt(n()),
n_fd = n())
# Combine the two separate data frames.
sd_fd_summary <- left_join(stabiliation_summary, fitdiff_summary)
# Some of the FDs are negative, let's flip these to be positive
# and also flip the label so that the first species in the name
# is always the fitness superior.
sd_fd_summary <- sd_fd_summary |>
# if mean_fd is < 0, the following command gets the absolute
# value and also flips around the species code so that
# the fitness superior is always the first species in the code
mutate(pair = ifelse(mean_fd < 0,
paste0(str_extract(pair, "..$"),
"_",
str_extract(pair, "^..")),
pair),
mean_fd = abs(mean_fd))
sd_fd_summary <-
sd_fd_summary |>
mutate(
# outcome = ifelse(mean_fd - 2*sem_fd >
# mean_sd + 2*sem_sd, "exclusion", "neutral"),
outcome2 = ifelse(mean_fd > mean_sd, "exclusion", "coexistence"),
outcome2 = ifelse(mean_sd < 0, "exclusion or priority effect", outcome2)
)
base_w_rug <-
base +
geom_point(data = sd_fd_summary,
aes(x = mean_sd, y = 0), shape = "|", size = 5,
inherit.aes = F) +
theme(axis.title = element_blank())
base_w_rug
# Example point, using UR_FE values
base_w_rug +
annotate("pointrange", x = 0.275, y = 0.0591, ymin = 0.0591-0.307,ymax = 0.0591+0.307, shape = 21, stroke = 1.5, size = 1, linewidth = 0.1, fill = "#4477aa") +
annotate("pointrange", x = 0.275, y = 0.0591, xmin = 0.275-0.152,xmax = 0.275+0.152, shape = 21, fill = "#4477aa", linewidth = 0.1, size = 1, stroke = 1.5) +
annotate("text", x = 0.275 + 0.3, y = 0.0591 + 0.15, color = "#4477aa", label = "Festuca - Plantago", size = 8, fontface = "italic") 
base +
geom_pointrange(data = sd_fd_summary,
aes(x = mean_sd, y = mean_fd,
ymin = mean_fd-sem_fd,
ymax = mean_fd+sem_fd, fill = outcome2), size = 1, linewidth = 0.1,
shape = 21, stroke = 1.5) +
geom_pointrange(data = sd_fd_summary,
aes(x = mean_sd, y = mean_fd,
xmin = mean_sd-sem_sd,
xmax = mean_sd+sem_sd, fill = outcome2), size = 1,
linewidth = 0.1, shape = 21, stroke = 1.5) +
scale_fill_manual(values = c("#4477aa", "#ee6677", "#ccbb44")) +
theme(legend.position = "none")+
theme(axis.title = element_blank())
Detailed in Kandlikar et al. (2021)
In California grasslands, microbes tend to drive stronger fitness differences than stabilization.
Can we answer this question more generally?

Data for 518 pairwise comparisons of stabilization and fitness differences (results shown for 72 pairs here)
Results in Yan, Levine, and Kandlikar (2022); Analyses available on Zenodo
Microbially-mediated coexistence in pine-oak forests in Spain, Pajares-Murgó et al. (2024)
Unceded Chumash territory
2. Microbial effects on plant communities in variable environments
General themes:
Advancing ecological insights in light of conservation and management challenges
Situating our understanding in historical contexts
library(sf)
library(tidyverse)
little <- sf::st_read("../img/pinupalu.shp")
## Reading layer `pinupalu' from data source
## `/home/gkandlikar@lsu.edu/gklab/talks/img/pinupalu.shp' using driver `ESRI Shapefile'
## Simple feature collection with 38 features and 5 fields
## Geometry type: POLYGON
## Dimension: XY
## Bounding box: xmin: -95.21806 ymin: 26.61921 xmax: -75.79999 ymax: 36.84856
## CRS: NA
# States we want to show
llp_states <- c("texas", "louisiana","mississippi", "alabama", "florida", "georgia", "south carolina", "north carolina", "virginia", "tennessee")
states <- map_data("state") |>
filter(region %in% llp_states)
ggplot() +
geom_polygon(data = states,
aes(x = long, y = lat, group = group), fill = "transparent", color = 'darkgrey') +
geom_sf(data = little, fill = alpha('#228833', 0.5)) +
theme_void()
Turpentine Industry from Frank Leslie’s Illustrated Newspaper. Link.
Photo from Longleaf Alliance. Link.
Photo: Kandlikar lab members at Lee Memorial Forest (LSU Ag Center), Franklinton LA
Prescribed burn at LSU Forestry Camp, 13 April 2026
Still image from 1920s US Forest Service film “When Trees Talk”. Video available here.


Remarkable diversity at small spatial scales (40-50 plant spp/m2)
Woody encroachment pervasive under fire suppression
Over 500 million hectares of grasslands/savannas affected
Many drivers…
Over 500 million hectares of grasslands/savannas affected
Many drivers… but we wonder if there is a role for plant–soil microbial feedbacks.

Two lines of evidence.
# curl::curl_download("https://datadryad.org/downloads/file_stream/2789266", destfile = "../data/jiang2024-dataset.csv")
jiang_dat <- read_csv("../data/jiang2024-dataset.csv")
# jiang_metamod <- metafor::rma.mv(rr, var, data = jiang_dat, mods = ~ Life.form-1)
# saveRDS(jiang_metamod, "../data/jiang-metamod.rds")
metamod <- readRDS("../data/jiang-metamod.rds")
metamod_s <- broom::tidy(metamod)
jiang_plot <-
jiang_dat |>
ggplot(aes(x = rr, y = Life.form, color = Life.form)) +
ggbeeswarm::geom_quasirandom(shape = 21) +
scale_color_manual(values = c("#ccbb44","transparent")) +
annotate("point",
x = metamod_s$estimate[1], y = 1, size = 3, shape = 21, stroke = 1.5) +
# annotate("point",
# x = metamod_s$estimate[2], y = 2, size = 3, shape = 21, stroke = 1.5) +
ylab("") +
xlab("Growth in self vs. growth in non-self") +
xlim(-2.5,2.5)+
geom_vline(xintercept = 0, linetype = 'dashed') +
theme(legend.position = "none",
axis.text.y = element_text(size = 12, color = 'black'))
jiang_plot
Data from Jiang et al. (2024)

Data from Jiang et al. (2024)
In African savannas, N-fixing legumes are the primary woody invader in >90% of studies sites (Stevens et al. 2017)
Ectomycorrhizal symbiosis is especially more common among encroaching woody plants (Simha and Kandlikar 2026)
# download.file("https://www.science.org/doi/suppl/10.1126/science.aai8212/suppl_file/bennett_aai8212_database-s1.xlsx", destfile = "../data/bennett2017-dataset.xlsx")
readxl::read_xlsx("../data/bennett-2017.xlsx", sheet = 2) |>
mutate(`Type of Mycorrhiza` = factor(`Type of Mycorrhiza`, c("EM", "AM"))) |>
mutate(logratio = log(`Average of Biomass in Conspecific Soil`/`Average of Biomass in Heterospecific Soil`)) |>
arrange(logratio) |>
mutate(rown = row_number()) |>
ggplot(aes(x = logratio, color = `Type of Mycorrhiza`, y = rown, shape = `Type of Mycorrhiza`)) +
geom_point(size = 2) +
geom_segment(aes(x = 0, xend = logratio), linewidth = 0.125) +
xlab("Growth in self-conditioned microbes vs.\n nonself-conditioned microbes") +
geom_vline(xintercept = 0) +
scale_color_manual(values = c("#0077bb", "#cc3311")) +
theme(axis.line.y = element_blank(),
axis.text.y = element_blank(),
axis.title.y = element_blank(),
axis.ticks.y = element_blank(),
legend.position = "inside",
legend.position.inside = c(0.9, 0.1),
legend.text = element_text(size = 12),
legend.title = element_blank())

Data from Bennett et al. (2017)
Approach:
Mathematical modeling: experimentally-informed dynamics of plant community dynamics
Field observations and experiment: evaluate fire effects on soil communities
Plant growth experiment: evaluate shrub and grass growth in pre- and post-fire shrub- and grass-conditioned soils
Biological aspects to consider:
Grasses and woody plants can have distinct microbial feedbacks.
Microbial legacies develop over time, and may develop faster or slower for grasses vs. woody plants.
Grasses and woody plants respond differently to fire.
Woody adults and woody seedlings respond differently to fire.
Fires can vary in their intensity and their frequency.
Microbes can affect various aspects of demography, but as a starting point we assume they regulate early life success (germination and seedling survival)
\[ \frac{dP_{00}}{dt} = - \overbrace{r_s P_{00}(P_{ss})}^{\substack{\text{woody} \\ \text{establishment}}}- \overbrace{r_g P_{00}(P_{{g0}+}P_{gg}+P_{gs})}^{\text{grass establishment}} + \overbrace{\mu_g P_{g0} +\mu_\sigma P_{\sigma 0}}^{\substack{\text{mortality in} \\ \text{unconditioned patch}}} + \overbrace{d_g P_{0g} + d_s P_{0s}}^{\substack{\text{microbial decay in} \\ \text{unoccupied patch}}} \qquad(1)\]
\[ \frac{dP_{g0}}{dt} = \overbrace{r_g P_{00}(P_{g0} + P_{gg} + P_{gs})}^{\substack{\text{grass establishment in} \\ \text{unconditioned patch}}} - \overbrace{\mu_g P_{g0}}^{\substack{\text{grass} \\ \text{mortality}}} - \overbrace{c_g P_{g0}} ^{\substack{\text{grass} \\ \text{conditioning}}} \qquad(2)\]
\[ \frac{dP_{\sigma 0}}{dt} = \overbrace{r_s P_{00}(P_{ss})}^{\substack{\text{woody establishment in} \\ \text{unconditioned patch}}} - \overbrace{\mu_\sigma P_{\sigma 0}}^{\substack{\text{woody} \\ \text{mortality}}} - \overbrace{c_s P_{\sigma 0}}^{\substack{\text{woody} \\ \text{conditioning}}} \qquad(3)\]\[ \frac{dP_{gg}}{dt} = \overbrace{r_g m_{gg} P_{0g}(P_{g0} + P_{gg} + P_{gs})}^{\substack{\text{grass establishment in} \\ \text{grass-conditioned patch}}} + \overbrace{c_g (P_{g0} + P_{gs})}^{\text{conditioning}} - \overbrace{\mu_g P_{gg}}^{\text{mortality}} \qquad(4)\]
\[ \frac{dP_{\sigma s}}{dt} = \overbrace{r_s m_{ss} P_{0s}P_{ss}}^{\substack{\text{woody establishment in} \\ \text{woody-conditioned patch}}} - \overbrace{c_s P_{\sigma s}}^{\substack{\text{growth into} \\ \text{adult stage}}} - \overbrace{\mu_\sigma P_{\sigma s}}^{\text{mortality}} \qquad(5)\]
\[ \frac{dP_{ss}}{dt} = \overbrace{c_s (P_{\sigma 0} + P_{\sigma g})}^{\substack{\text{conditioning by and growth of}\\ \text{woody seedlings} }} + \overbrace{c_s (P_{\sigma s})}^{\substack{\text{ growth of}\\ \text{woody seedlings} }} - \overbrace{\mu_s P_{ss}}^{\text{mortality}} \qquad(6)\]
\[ \frac{dP_{gs}}{dt} = \overbrace{r_g m_{gs} P_{0s} (P_{g0} + P_{gg} + P_{gs})}^{\substack {\text{grass establishment in} \\ \text{woody-conditioned patch}}} - \overbrace{\mu_g P_{gs}}^{\text{mortality}} - \overbrace{c_g P_{gs}}^{\text{conditioning}} \qquad(7)\]
\[ \frac{dP_{\sigma g}}{dt} = \overbrace{r_s m_{sg} P_{0g} (P_{ss})}^{\substack {\text{woody establishment in} \\ \text{grass-conditioned patch}}} - \overbrace{\mu_\sigma P_{\sigma g}}^{\text{mortality}} - \overbrace{c_s P_{\sigma g}}^{\text{conditioning}} \qquad(8)\]\[ \frac{dP_{0g}}{dt} = - \overbrace{r_g m_{gg} P_{0g}(P_{g0}+P_{gg}+P_{gs})}^{\text{grass establishment}} - \overbrace{r_s m_{sg} P_{0g}(P_{ss})}^{\substack{\text{woody} \\ \text{establishment}}} + \overbrace{\mu_g P_{gg} + \mu_\sigma P_{\sigma g}}^{\substack{\text{mortality in} \\ \text{grass-conditioned patch}}} - \overbrace{d_g P_{0g}}^{\substack{\text{microbial} \\ \text{decay}}} \qquad(9)\]
\[ \begin{multline} \frac{dP_{0s}}{dt} = - \overbrace{r_s m_{ss} P_{0s}(P_{ss})}^{\text{woody establishment}} - \overbrace{r_g m_{gs} P_{0s}(P_{g0}+P_{gg}+P_{gs})}^{\text{grass establishment}} + \\ \overbrace{\mu_s P_{ss} + \mu_\sigma P_{\sigma s} + \mu_g P_{gs}}^{\substack{\text{mortality in} \\ \text{woody-conditioned patch}}} - \overbrace{d_s P_{0s}}^{\substack{\text{microbial} \\ \text{decay}}} \end{multline} \qquad(10)\]library(tidybayes)
meta <- readRDS("../img/swj-brms-meta.RDS")
b_draws <-
meta |>
spread_draws(b_Life.formNonwoody, b_Life.formWoody) |>
rename(mgg_draws = b_Life.formNonwoody,
mss_draws = b_Life.formWoody) |>
median_qi() |>
mutate(across(is.numeric, exp))
plot_det_pt <-
plot_det +
geom_pointrange(inherit.aes = F,
data = b_draws,
aes(x = mgg_draws, y = mss_draws,
xmin = mgg_draws.lower, xmax = mgg_draws.upper),
color = 'white', shape = 21, size = 1, stroke = 2) +
geom_pointrange(inherit.aes = F,
data = b_draws,
aes(x = mgg_draws, y = mss_draws,
ymin = mss_draws.lower, ymax = mss_draws.upper),
color = 'white', shape = 21, size = 1, stroke = 2) +
labs(caption = "White point & error bars show median & 95 CrI of Bayesian meta-analysis\nof grasses and woody plants from areas undergoing encroachment.")
plot_det_pt
Fires of higher intensities cause more mortality of grasses, woody seedlings, and soil microbes.
In general, woody plants are more fire-sensitive than grasses, but their sensitivity varies by life stage.
Three scenarios of fire impacts on woody adults:
int = seq(0,1, length.out = 20)
fire = data.frame(int = seq(0,1, length.out = 20),
grass = 1/(1+150*exp(-10.02*int)),
seedling = 1/(1+150*exp(-20.04*int)),
adult = 1/(1+150*exp(-10.02*int)),
sens = rep("Fire Sensitive", 20)) |>
pivot_longer(cols=c(grass,seedling,adult), names_to = "Plant", values_to = "mort")
fire2 = fire |> mutate(sens = "Less Sensitive",
mort = case_when(Plant == "adult" ~ mort/2, TRUE ~ mort))
fire3 = fire |> mutate(sens = "Not Sensitive",
mort = case_when(Plant == "adult" ~ 0, TRUE ~ mort))
fire_all = rbind(fire, fire2, fire3) |> mutate(sens = as.factor(sens), Plant = as.factor(Plant))
levels(fire_all$sens)
## [1] "Fire Sensitive" "Less Sensitive" "Not Sensitive"
levels(fire_all$Plant) = c("Woody Adult", "Grass", "Woody Seedling")
fire_all$Plant = factor(fire_all$Plant, levels = c("Grass", "Woody Seedling", "Woody Adult"))
scenarios = ggplot(data = fire_all, aes(x = int, y = mort, group = Plant)) +
geom_path(aes(color = Plant, linetype = Plant), linewidth = 3) +
scale_color_manual(values = c("#FDE725FF", "#2A788EFF", "#440154FF"))+
scale_linetype_manual(values = c("solid", "dashed", "longdash"))+
labs(x = "Fire Intensity", y="Percent Mortality")+
scale_x_continuous(expand = c(0,0), breaks = c(0.25, 0.5, 0.75, 1.0)) +
scale_y_continuous(expand = c(0,0))+
facet_wrap(~ sens)
scenarios



Interpretation:
Microbial feedbacks constrain which fire regimes sustain grassy communities
Microbial feedbacks can also constrain fire-based grassland restoration after woody encroachment
Detailed in preprint (Simha and Kandlikar 2026), feedback welcome
Approach:
1. Field observations and experiment: evaluate fire effects on soil communities
1. Plant growth experiment: evaluate shrub and grass growth in pre- and post-fire shrub- and grass-conditioned soils


Working in collaboration with the Southeastern Tribal Alliance for Fire (SETA Fire), we hope to…
Ecological theory for microbial effects on plant coexistence
Microbial effects on plant communities in variable environments
Theme 1: Microbial contributions to plant eco–evolutionary dynamics
brapa <-
read_csv("brapa-data.csv") |>
pivot_wider(names_from = Type, values_from = y) |>
mutate(Seed = ifelse(Seed == "HW", "Control", "Low Water"),
Soil = ifelse(Soil == "HW", "Control", "Low Water"),
Current = ifelse(Current == "HW", "Control", "Low Water"))
brapa |>
group_by(Current) |>
summarize(mean = mean(mean)) |>
ggplot(aes(x = Current, y = mean)) +
geom_point(size = 4) +
ylim(0,250) +
labs(caption = "Marginal means from a Bayesian GLM (Poisson family)",
y = "Per-capita fecundity",
xlab = "")
brapa |>
ggplot(aes(x = Current, y = mean, ymin = sdlo, ymax = sdhi,
color = Soil, shape = Seed)) +
geom_pointrange(size = 1, position = position_dodge2(width = 0.3)) +
scale_color_manual(values = c("#4477aa", "#ee6677")) +
ylim(0,250) +
labs(caption = "Marginal means & 95% CrIs from a Bayesian GLM (Poisson family)",
y = "Per-capita fecundity",
xlab = "")
Ongoing efforts to uncover the mechanistic basis.

Theme 2: Microbial impacts on tropical forest communities
Focus area 1: Disruptions to plant–soil feedback under habitat fragmentation (Saini et al. accepted at AJB)

Focus area 2: Plant–fungal–nutrient interactions as drivers of monodominance of ectomycorrhizal-associating trees in hyperdiverse forests

Ecological theory for microbial effects on plant coexistence
Microbial effects on plant communities in variable environments



Thank you!
slides: https://talks.gklab.org/utep-26
contact: gkandlikar@lsu.edu