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Assessing seed and microsite limitation on population dynamics of a gypsophyte through experimental soil crust disturbance and seed addition

Matthew R. Tye1,4,Merari E. Ferrer-Cervantes1, Ana M. Sánchez3, Ana I. García-Cervigón2, Adrián Escudero3, María J. Albert3, José M. Olano2, José M. Iriondo3, Pedro F. Quintana-Ascencio1*

1 Department of Biology, University of Central Florida, Orlando, FL 32816, US

2 Dpto. de Ciencias Agroforestales, Escuela de Ingenierías Agrarias, Universidad de Valladolid, Los Pajaritos s/n. 42003 Soria, Spain

3 Área de Biodiversidad y Conservación, Universidad Rey Juan Carlos, Móstoles, 28040 Madrid, Spain

4 Department of Plant Ecology and Evolution, Uppsala University, Uppsala, Sweden 75236

*

Figure 1. Histogram showing average June to September temperatures of our three study years (red lines) in comparion to historic data from 1976 to 2012 recorded at the Aranjuez meteorological station (data provided by the Spanish Meteorological Agency AEMET,

Figure2. Histogram showing average January to May precipiation of our three study years (red lines) in comparion to historic data from 1976 to 2012 recorded at the Aranjuez meteorological station (data provided by the Spanish Meteorological Agency AEMET,

Figure 3. Overall IPM kernels for site 1 2008-09 (A), 2009-10 (B) and 2010-11 (C) and site 2 2008(D), 2009(E) and 2010(F). These diagrams are representations of the population growth models (as explained in Metcalf et al. 2013). Each matrix has 201 by 201 cells. The diagonal region increasing from left to right corresponds to survival and growth. The region at the bottom-right corresponds to the seeds being produced that directly germinate within the span of the observed annual transition, thus not entering the persistent seedbank. The thin, white line at the extreme bottom of the graph corresponds to seeds entering the seedbank. Lighter colors represent higher relative magnitude for the focal transition.

Table 1. Plot length (wide was 1 m for all plots), and densities of adults and seedling at each plot per site split by treatment and by year.

Site / plot / Plot longitude (m) / Treatment / Plants/m2 / Seedlings/m2
2009 / 2010 / 2011 / 2009 / 2010 / 2011
1 / 1 / 5.4 / control / 19.6 / 7.6 / 2.0 / 175.0 / 175.2 / 18.9
1 / 6 / 4.4 / control / 11.6 / 7.7 / 3.4 / 346.8 / 215.5 / 31.6
1 / 14 / 2.4 / control / 21.3 / 15.0 / 4.2 / 62.1 / 125.8 / 25.4
1 / 2 / 2.6 / disturbance / 49.6 / 31.9 / 13.5 / 266.2 / 252.7 / 46.5
1 / 4 / 1.8 / disturbance / 62.2 / 34.4 / 14.4 / 671.7 / 286.1 / 72.2
1 / 7 / 3.2 / disturbance / 35.0 / 21.3 / 4.7 / 356.3 / 229.4 / 50.6
1 / 3 / 3.4 / seed addition / 33.5 / 21.2 / 8.2 / 793.2 / 506.2 / 121.2
1 / 5 / 3.0 / seed addition / 40.0 / 25.7 / 5.3 / 398.7 / 347.0 / 88.7
1 / 15 / 1.6 / seed addition / 48.1 / 16.9 / 5.6 / 71.3 / 128.1 / 35.0
2 / 9 / 4.0 / control / 25.8 / 13.0 / 3.3 / 437.5 / 15.9 / 12.5
2 / 12 / 4.4 / control / 23.0 / 9.8 / 2.5 / 105.2 / 37.0 / 6.6
2 / 8 / 3.2 / disturbance / 33.8 / 21.3 / 4.7 / 584.4 / 61.9 / 9.7
2 / 13 / 6.2 / disturbance / 16.0 / 9.8 / 3.4 / 81.9 / 17.7 / 8.1
2 / 10 / 7.4 / seed addition / 13.5 / 9.2 / 3.8 / 207.6 / 114.9 / 25.1
2 / 11 / 3.2 / seed addition / 37.2 / 17.8 / 4.1 / 332.7 / 168.4 / 47.2
/ 35.7 / 20.2 / 6.8 / 349.0 / 251.8 / 54.5
24.9 / 13.5 / 3.6 / 291.6 / 69.3 / 18.2

Table 2. AIC tables showing differing general linear models for survival (A), probability of reproduction (B) and seeds per plant (C).

A.GLM for survival (all years)

Models for survival / K / AICc / Delta_AICc
Survival ~ log-diameter + logdiameter^2 + Treat* Sites + TJS + PJM / 9 / 3353.3 / 0.0
Survival ~ log-diameter + logdiameter^2 + Treat + Sites + TJS + PJM / 7 / 3355.4 / 2.1
Survival ~ log-diameter + logdiameter^2 + Treat* Sites + TJS / 8 / 3357.7 / 4.4
Survival ~ log-diameter + logdiameter^2 + Treat* Sites *TJS*PJM / 19 / 3359.5 / 6.2
Survival ~ log-diameter + logdiameter^2 + Treat + TJS / 5 / 3360.1 / 6.8
Survival ~ log-diameter + logdiameter^2 + Treat + Sites + TJS / 6 / 3360.8 / 7.5
Survival ~ log-diameter + logdiameter^2 + Treat*TJS / 7 / 3362.7 / 9.4
Survival ~ log-diameter + logdiameter^2 + Treat* Sites *TJS / 13 / 3362.8 / 9.5
Survival ~ log-diameter + logdiameter^2 + TJS / 3 / 3403.1 / 49.8
Survival ~ log-diameter + logdiameter^2 + Treat + PJM / 5 / 3408.0 / 54.7
Survival ~ log-diameter + logdiameter^2 + Treat + Sites + PJM / 6 / 3409.5 / 56.2
Survival ~ log-diameter + logdiameter^2 + Treat* Sites + PJM / 8 / 3410.1 / 56.8
Survival ~ log-diameter + logdiameter^2 + Treat*PJM / 7 / 3411.1 / 57.8
Survival ~ log-diameter + logdiameter^2 + Treat* Sites *PJM / 13 / 3413.7 / 60.4
Survival ~ log-diameter + logdiameter^2 + PJM / 3 / 3432.0 / 78.7
Survival ~ log-diameter + logdiameter^2 + Treat / 4 / 3569.9 / 216.6
Survival ~ log-diameter + logdiameter^2 + Treat + Sites / 5 / 3571.3 / 218.0
Survival ~ log-diameter + logdiameter^2 + Treat* Sites / 7 / 3571.8 / 218.5
Survival ~ log-diameter + logdiameter^2 / 2 / 3589.2 / 235.9
Survival ~ log-diameter + logdiameter^2 + Sites / 3 / 3589.9 / 236.6
Survival ~ 1 / 1 / 3628.5 / 275.2
Treat= treatments, TJS= temperature June-September, PJM= precipitation January-May

Summary for the most informative model for survival

Glm (formula = survival ~ log-diameter + logdiameter^2+ Treat * Sites + TJS + PJM, family = binomial)
Coefficients: / Estimate / Std. Error / z value / Pr(>|z|)
(Intercept) / 12.54 / 2.54 / 4.93 / 0.00 / ***
Log-diameter / 3.08 / 0.39 / 7.96 / 0.00 / ***
logsquare / -0.53 / 0.09 / -5.96 / 0.00 / ***
Treat 0-1 / 0.96 / 0.15 / 6.32 / 0.00 / ***
Treat 0-2 / 0.48 / 0.15 / 3.27 / 0.00 / **
Sites 1-2 / 0.02 / 0.16 / 0.14 / 0.89
TJS / -0.76 / 0.10 / -7.21 / 0.00 / ***
PJM / 0.01 / 0.001 / 3.35 / 0.00 / ***
Treat 0-1:Site 1-2 / -0.47 / 0.22 / -2.15 / 0.03 / *
Treat 0-1:Site 1-2 / 0.04 / 0.22 / 0.18 / 0.85
Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’
Models for probability of reproduction / K / AICc / Delta_AICc
Rep ~ log-diameter + Treat*Sites + TJS + PJM / 9 / 1186.4 / 0.0
Rep ~ log-diameter + Treat + Sites + TJS + PJM / 7 / 1202.8 / 16.3
Rep ~ log-diameter + Treat*Sites*PJM / 13 / 1276.1 / 89.6
Rep ~ log-diameter + Treat*Sites + PJM / 8 / 1306.7 / 120.3
Rep ~ log-diameter + Treat + Sites + PJM / 6 / 1322.1 / 135.6
Rep ~ log-diameter + Treat*PJM / 7 / 1364.5 / 178.0
Rep ~ log-diameter + PJM / 3 / 1378.8 / 192.4
Rep ~ log-diameter + Treat + PJM / 5 / 1379.8 / 193.4
Rep ~ log-diameter + Treat*Sites*TJS / 13 / 1939.0 / 752.6
Rep ~ log-diameter + Treat*TJS / 7 / 2032.2 / 845.7
Rep ~ log-diameter + Treat*Sites + TJS / 8 / 2035.8 / 849.4
Rep ~ log-diameter + Treat + Sites + TJS / 6 / 2046.7 / 860.2
Rep ~ log-diameter + TJS / 3 / 2077.2 / 890.8
Rep ~ log-diameter + Treat + TJS / 5 / 2077.9 / 891.4
Rep ~ log-diameter + Treat*Sites / 7 / 3140.6 / 1954.2
Rep ~ log-diameter + Treat + Sites / 5 / 3143.7 / 1957.2
Rep ~ log-diameter + Sites / 3 / 3148.6 / 1962.2
Rep ~ log-diameter + Treat / 4 / 3164.8 / 1978.4
Rep ~ log-diameter / 2 / 3171.5 / 1985.0
Rep ~ 1 / 1 / 3799.3 / 2612.9
Treat= treatments, TJS= temperature June-September, PJM= precipitation January-May

B.GLM’s for probability of reproduction (all years)

Summary for the most informative model for probability of reproduction

Glm (formula = rep ~ logdiameter + Treat*Sites + TJS + PJM, family = binomial)
Estimate / Std. Error / z value / Pr(>|z|)
(Intercept) / -251.34 / 23.58 / -10.66 / < 2e-16 / ***
Log-diameter / 5.30 / 0.27 / 19.64 / < 2e-16 / ***
Treat 0-1 / -0.57 / 0.28 / -2.06 / 0.04 / *
Treat 0-2 / -1.10 / 0.28 / -3.99 / 0.00 / ***
Sites 1-2 / 0.41 / 0.29 / 1.41 / 0.16
TJS / 8.78 / 0.91 / 9.66 / < 2e-16 / ***
PJM / 0.20 / 0.01 / 14.23 / < 2e-16 / ***
Treat 0-1:Sites 1-2 / 0.47 / 0.39 / 1.22 / 0.22
Treat 0-2:Sites 1-2 / 1.60 / 0.38 / 4.21 / 0.00 / ***
Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’

C.GLM’s for seeds per plant (all years)

Model for seeds per plant / K / AICc / Delta_AICc
Log_seed_plant ~ log-diameter + Treat*Sites + TJS + PJM / 10 / 2593.0 / 0.0
Log_seed_plant ~ log-diameter + Treat + Sites + TJS + PJM / 8 / 2599.7 / 6.7
Log_seed_plant ~ log-diameter + Treat* Sites + TJS / 9 / 2616.1 / 23.1
Log_seed_plant ~ log-diameter + Treat + Sites + TJS / 7 / 2622.7 / 29.7
Log_seed_plant ~ log-diameter + Treat*TJS / 8 / 2772.1 / 179.1
Log_seed_plant ~ log-diameter + Treat + TJS / 6 / 2776.3 / 183.3
Log_seed_plant ~ log-diameter + TJS / 4 / 2803.5 / 210.4
Log_seed_plant ~ log-diameter + Treat* Sites + PJM / 9 / 2811.5 / 218.5
Log_seed_plant ~ log-diameter + Treat + Sites + PJM / 7 / 2816.7 / 223.7
Log_seed_plant ~ log-diameter + Treat* Sites / 8 / 2837.3 / 244.3
Log_seed_plant ~ log-diameter + Treat + Sites / 6 / 2842.1 / 249.1
Log_seed_plant ~ log-diameter + Sites / 4 / 2853.5 / 260.5
Log_seed_plant ~ log-diameter + Treat*PJM / 8 / 2935.5 / 342.5
Log_seed_plant ~ log-diameter + Treat + PJM / 6 / 2952.0 / 359.0
Log_seed_plant ~ log-diameter + PJM / 4 / 2959.0 / 366.0
Log_seed_plant ~ log-diameter + Treat / 5 / 2979.2 / 386.2
Log_seed_plant ~ log-diameter / 3 / 2985.7 / 392.7
Log_seed_plant ~ 1 / 2 / 4602.1 / 2009.1

Summary for the most informative model for seeds per plant

Glm (log_seed_plant ~ log-diameter + Treat*Sites + TJS + PJM, family=gaussian)
Coefficients: / Estimate / Std. Error / t value / Pr(>|t|)
(Intercept) / -16.95 / 1.07 / -15.90 / < 2e-16 / ***
Log-diameter / 2.00 / 0.04 / 48.00 / < 2e-16 / ***
Treat 0-1 / -0.38 / 0.07 / -5.32 / 0.00 / ***
Treat 0-2 / -0.18 / 0.07 / -2.54 / 0.01 / *
Sites 1-2 / -0.72 / 0.07 / -9.79 / < 2e-16 / ***
TJS / 0.75 / 0.05 / 15.50 / < 2e-16 / ***
PJM / 0.00 / 0.00 / 5.02 / 0.00 / ***
Treat 0-1:Sites 1-2 / 0.18 / 0.10 / 1.80 / 0.07 / .
Treat 0-2:Sites 1-2 / 0.33 / 0.10 / 3.27 / 0.00 / **
Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’

D) Summary for the most informative model for plant growth

Glm (log-diameter (t+1) ~ log-diameter + Treat*Sites + TJS + PEM, family=gaussian)

Coefficients: / Estimate / Std. Error / t value / Pr(>|t|)
(Intercept) / -4.0152 / 0.446959 / -8.983 / < 2e-16 ***
Log-diameter / 0.779462 / 0.016123 / 48.346 / < 2e-16 ***
Treat 0-1 / -0.14482 / 0.030441 / -4.757 / 2.19e-06 ***
Treat 0-2 / -0.11433 / 0.030762 / -3.717 / 0.000211 ***
Sites 1-2 / -0.07377 / 0.035276 / -2.091 / 0.036693 *
TJS / 0.216194 / 0.018852 / 11.468 / < 2e-16 ***
PEM / -0.00076 / 0.000209 / -3.657 / 0.000266 ***
Treat 0-1:Sites 1-2 / 0.120727 / 0.045445 / 2.657 / 0.007994 **
Treat 0-1:Sites 1-2 / 0.117657 / 0.045499 / 2.586 / 0.009823 **

Table 3. Summary of a linear model of the effect of year, site and treatment on number of seedlings m-2 (logarithm transformed; R2=0.72, F = 20.36 on 5 and 39 degrees of freedom, p <0.001). This model was chosen because parsimony since a model with interactive effects provided similar evidence (AICc weight < 0.5)

Estimate / Std. Error / t value / Pr(>|t|)
Intercept / 5.44 / 0.25 / 21.49 / <0.001
Year: 2010 / -0.70 / 0.26 / -2.69 / 0.01
Year: 2011 / -2.15 / 0.26 / -8.27 / <0.001
Disturbance / 0.47 / 0.26 / 1.82 / 0.077
Seed addition / 0.87 / 0.26 / 3.35 / 0.002
Site2 / -0.96 / 0.22 / -4.40 / <0.001