Data from: Development time and host-parasitoid stability: An experimental test
Data files
Mar 31, 2026 version files 334.98 KB
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Bean_fate_LD-SD_experiment.csv
80.51 KB
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Control_simulation_model_ndnp.R
20.02 KB
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Control_simulation_model.R
23.94 KB
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Host_fate.R
5.80 KB
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Host_feeding.csv
4.88 KB
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Host_feeding.R
4.28 KB
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LD_simulation_model_ndnp.R
21.95 KB
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LD_simulation_model.R
26.86 KB
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LD-SD_experiment_time_series_block1.csv
26.36 KB
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LD-SD_experiment_time_series_block2.csv
13.74 KB
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power-period_repeated_measures.csv
5.82 KB
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Power-period_repeated_measures.R
12.94 KB
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README.md
22.63 KB
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SD_simulation_model_days_ndnp.R
20.04 KB
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SD_simulation_model_days.R
24.41 KB
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Time_series_analysis2.R
18.85 KB
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treatment_mean-SD_log10.csv
1.95 KB
Apr 08, 2026 version files 330.84 KB
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Bean_fate_LD-SD_experiment.csv
80.51 KB
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Control_simulation_model_ndnp2.R
20.01 KB
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Control_simulation_model2.R
23.87 KB
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Host_fate.R
5.80 KB
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Host_feeding.csv
4.88 KB
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Host_feeding.R
4.28 KB
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LD_simulation_model_ndnp2.R
21.94 KB
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LD_simulation_model2.R
26.84 KB
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LD-SD_experiment_time_series_block1.csv
26.36 KB
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LD-SD_experiment_time_series_block2.csv
13.74 KB
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power-period_repeated_measures.csv
5.82 KB
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Power-period_repeated_measures.R
12.94 KB
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README.md
22.58 KB
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SD_simulation_model_ndnp2.R
18.70 KB
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SD_simulation_model2.R
21.77 KB
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Time_series_analysis2.R
18.85 KB
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treatment_mean-SD_log10.csv
1.95 KB
Abstract
The duration of specific life stages and relative lengths of the predator and prey life cycles can, in theory, exert strong influences on population dynamics and may be important considerations in managing pest outbreaks. However, rigorous experimental tests of this theory do not exist. We conducted a microcosm experiment using the cowpea weevil (Callosobruchus maculatus) and its parasitoid (Anisopteromalus calandrae) to assess how a 60% increase (long-duration treatment) or 60% decrease (short-duration treatment) in the invulnerable (late larval to mid pupal) host stage affected host-parasitoid population dynamics. We predicted that stability is most likely for microcosms in the short-duration treatment, whereas generation cycles should occur for the long-duration and control treatments, with a longer period in the long-duration treatment. Our results largely support these predictions. In the control treatment, persistent oscillations occurred with a period of about one generation, consistent with theory. The long-duration treatment produced cycles similar in period and strength to the control, rather than predicted longer period ones. As expected, cycles disappeared in the short-duration treatment, and host densities were suppressed by 41-49% relative to the other treatments. Variability in host densities in the short-duration treatment was high, likely a consequence of demographic stochasticity in small populations, whereas variability in parasitoid densities was 24-27% lower than in the other treatments. Complementary simulation models that closely matched the biological details and protocols used in the experiment reproduced the observed dynamics in the short-duration and control treatments. The oscillatory period observed in the long-duration treatment was also replicated when stronger, scramble-type competition in the weevil attack process was incorporated into the model. We conclude that age structure and development time are critical determinants of host-parasitoid population dynamics and should be explicitly considered in the selection of biological control agents and in evaluations of program success or failure.
Dataset DOI: 10.5061/dryad.05qfttfj4
Description of the data and file structure
Data for all analyses are provided as *.csv files. Each csv file has an associated R code that was used for the analysis of these data. R version 4.3.1. Model simulations were also conducted using R. For these simulations, separate models and code were used for each of the three treatments (control, long-duration, and short-duration). Models for each treatment are also divided into those for a scenario with host beans added at 12-day intervals (pulsed addition) and a scenario with host beans added daily (no pulse). The latter R files have a “_ndnp2.R”.
Files and variables
File: Bean_fate_LD-SD_experiment.csv
Description: This dataset summarizes the experimental study on the fate of moth beans in the experimental microcosms. Codes for each column of data are as follows:
Variables
- Date: the date that the dish of beans was removed from the microcosm
- TRTMNT: the experimental treatment (CTRL = control, LD = long-duration treatment, SD = short-duration treatment)
- Dish: dish number collected during that time period
- Noegg: number of host eggs on the bean (out of 100)
- PrewinA: number of hosts alive in the pre-window (H2) life stage (out of 100)
- PrewinD: number of hosts dead in the pre-window (H2) life stage (out of 100)
- PostwinA: number of hosts alive in the post-window (H3) life stage (out of 100)
- PostwinD: number of hosts dead in the post-window (H3) life stage (out of 100)
- Wemerged: number of hosts that emerged as adults (out of 100)
- PlarvA: number of parasitoid larvae alive in beans
- PlarvD: number of parasitoid larvae dead in beans
- Pemerged: number of parasitoid adults that emerged from beans
- Unknown: number of beans where the fate is unknown
- PWnoemerge: proportion of beans where no host emerged (= (PrewinA+PostwinA)/(100-Noegg))
- Pnoegg: proportion of beans with no eggs (= Noegg/100)
- PPnoemerge: proportion of hosts that survived to the vulnerable host stage (H3) in which the parasitoid died and did not emerge as an adult (= PlarvA/(PlarvA+PlarvD+Pemerged))
- Ptism = proportion of hosts that were parasitized (= (PlarvA+PlarvD+Pemerged)/(PostwinA+Wemerged+ PlarvA+PlarvD+Pemerged+PostwinD))
File: Host_feeding.csv
Description: This dataset summarizes the experimental study on host feeding by Anisopteromalus calandrae. Codes for each column of data are as follows:
Variables
- Date: date trial was conducted
- Replicate: replicate number for the study for different combinations of the percentage of beans with vulnerable hosts and the number of parasitoids released.
- Pbean: the percentage of moth beans that contained a host suitable for attack by the parasitoid
- Hosts: number of beans infected by the host
- Ptoid: number of parasitoid females released into the microcosm to attack hosts
- Wemerged: number of adult weevils that developed to adults in the microcosm
- Wdeadh2: number of weevils that died in the H2 stage
- Wdeadh3: number of weevils that died in the H3 stage
- pDH2: proportion of weevils that died in the H2 stage (= Wdeadh2/Hosts)
- PtoidF: number of parasitoid offspring that developed into an adult female
- PtoidM: number of parasitoid offspring that developed into an adult male
- PCptized: per-capita number of hosts parasitized (=(PtoidF+PtoidM)/Ptoids)
- Ptism: proportion of hosts parasitized (=(PtoidF+PtoidM)/(Hosts-Wdeadh2))
- Pfeeding: proportion of hosts fed upon (= pDH2 – 0.0148); where 0.0148 is the mean for the Pbean Control microcosms for the proportion of hosts that die in the H2 stage (pDH2)
- Pcfeed: number of host fed upon per adult parasitoid released into the microcosm (= (Pfeeding*Hosts)/Ptoids)
- LPcfeed: ln(Pcfeed + 0.01)
- LoPfeed: ln(Pfeeding/(1-Pfeeding)+0.01)
File: LD-SD_experiment_time_series_block1.csv
Description: This dataset contains the raw time-series for host and parasitoid numbers per microcosm in the control, long-duration, and short-duration experimental treatments for the first temporal block. Codes for each column of data are as follows:
Variables
- census: census number
- Days: days since hosts were first released into each microcosm
- Date: date of census
- HaUC1: number of adult weevils alive at census t for Control treatment, replicate microcosm 1
- HdUC1: number of adult weevils dead at census t for Control treatment, replicate microcosm 1
- HsUC1: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 1
- PaUC1: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 1
- PdUC1: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 1
- PsUC1: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 1
- HaUC2: number of adult weevils alive at census t for Control treatment, replicate microcosm 2
- HdUC2: number of adult weevils dead at census t for Control treatment, replicate microcosm 2
- HsUC2: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 2
- PaUC2: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 2
- PdUC2: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 2
- PsUC2: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 2
- HaUC3: number of adult weevils alive at census t for Control treatment, replicate microcosm 3
- HdUC3: number of adult weevils dead at census t for Control treatment, replicate microcosm 3
- HsUC3: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 3
- PaUC3: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 3
- PdUC3: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 3
- PsUC3: total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 3
- HaUC4: number of adult weevils alive at census time for Control treatment, replicate microcosm 4
- HdUC4: number of adult weevils dead at census time for Control treatment, replicate microcosm 4
- HsUC4: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 4
- PaUC4: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 4
- PdUC4: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 4
- PsUC4: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 4
- HaUC5: number of adult weevils alive at census t for Control treatment, replicate microcosm 5
- HdUC5: number of adult weevils dead at census t for Control treatment, replicate microcosm 5
- HsUC5: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 5
- PaUC5: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 5
- PdUC5: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 5
- PsUC5: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 5
- HaLD1: number of adult weevils alive at census t for Long-duration treatment, replicate microcosm 1
- HdLD1: number of adult weevils dead at census t for Long-duration treatment, replicate microcosm 1
- HsLD1: sum total alive and dead adult weevils at census t for Long-duration treatment, replicate microcosm 1
- PaLD1: number of adult parasitoids alive at census t for Long-duration treatment, replicate microcosm 1
- PdLD1: number of adult parasitoids dead at census t for Long-duration treatment, replicate microcosm 1
- PsLD1: sum total alive and dead adult parasitoids at census t for Long-duration treatment, replicate microcosm 1
- HaLD5: number of adult weevils alive at census t for Long-duration treatment, replicate microcosm 5
- HdLD5: number of adult weevils dead at census t for Long-duration treatment, replicate microcosm 5
- HsLD5: sum total alive and dead adult weevils at census t for Long-duration treatment, replicate microcosm 5
- PaLD5: number of adult parasitoids alive at census t for Long-duration treatment, replicate microcosm 5
- PdLD5: number of adult parasitoids dead at census t for Long-duration treatment, replicate microcosm 5
- PsLD5: sum total alive and dead adult parasitoids at census t for Long-duration treatment, replicate microcosm 5
- HaSD1: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 1
- HdSD1: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 1
- HsSD1: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 1
- PaSD1: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 1
- PdSD1: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 1
- PsSD1: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 1
- HaSD2: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 2
- HdSD2: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 2
- HsSD2: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 2
- PaSD2: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 2
- PdSD2: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 2
- PsSD2: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 2
- HaSD3: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 3
- HdSD3: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 3
- HsSD3: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 3
- PaSD3: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 3
- PdSD3: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 3
- PsSD3: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 3
- HaSD4: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 4
- HdSD4: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 4
- HsSD4: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 4
- PaSD4: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 4
- PdSD4: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 4
- PsSD4: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 4
- HaSD5: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 5
- HdSD5: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 5
- HsSD5: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 5
- PaSD5: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 5
- PdSD5: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 5
- PsSD5: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 5
File: LD-SD_experiment_time_series_block2.csv
Description: This dataset contains the raw time-series for host and parasitoid numbers per microcosm in the control, long-duration, and short-duration experimental treatments for the second temporal block. Codes for each column of data are as follows:
Variables
- census: census number
- Days: days since hosts were first released into each microcosm
- Date: date of census
- HaUC1: number of adult weevils alive at census t for Control treatment, replicate microcosm 1
- HdUC1: number of adult weevils dead at census t for Control treatment, replicate microcosm 1
- HsUC1: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 1
- PaUC1: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 1
- PdUC1: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 1
- PsUC1: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 1
- HaUC3: number of adult weevils alive at census t for Control treatment, replicate microcosm 3
- HdUC3: number of adult weevils dead at census t for Control treatment, replicate microcosm 3
- HsUC3: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 3
- PaUC3: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 3
- PdUC3: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 3
- PsUC3: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 3
- HaUC4: number of adult weevils alive at census t for Control treatment, replicate microcosm 4
- HdUC4: number of adult weevils dead at census t for Control treatment, replicate microcosm 4
- HsUC4: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 4
- PaUC4: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 4
- PdUC4: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 4
- PsUC4: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 4
- HaUC5: number of adult weevils alive at census t for Control treatment, replicate microcosm 5
- HdUC5: number of adult weevils dead at census t for Control treatment, replicate microcosm 5
- HsUC5: sum total alive and dead adult weevils at census t for Control treatment, replicate microcosm 5
- PaUC5: number of adult parasitoids alive at census t for Control treatment, replicate microcosm 5
- PdUC5: number of adult parasitoids dead at census t for Control treatment, replicate microcosm 5
- PsUC5: sum total alive and dead adult parasitoids at census t for Control treatment, replicate microcosm 5
- HaLD1: number of adult weevils alive at census t for Long-duration treatment, replicate microcosm 1
- HdLD1: number of adult weevils dead at census t for Long-duration treatment, replicate microcosm 1
- HsLD1: sum total alive and dead adult weevils at census t for Long-duration treatment, replicate microcosm 1
- PaLD1: number of adult parasitoids alive at census t for Long-duration treatment, replicate microcosm 1
- PdLD1: number of adult parasitoids dead at census t for Long-duration treatment, replicate microcosm 1
- PsLD1: sum total alive and dead adult parasitoids at census t for Long-duration treatment, replicate microcosm 1
- HaLD3: number of adult weevils alive at census t for Long-duration treatment, replicate microcosm 3
- HdLD3: number of adult weevils dead at census t for Long-duration treatment, replicate microcosm 3
- HsLD3: sum total alive and dead adult weevils at census t for Long-duration treatment, replicate microcosm 3
- PaLD3: number of adult parasitoids alive at census t for Long-duration treatment, replicate microcosm 3
- PdLD3: number of adult parasitoids dead at census t for Long-duration treatment, replicate microcosm 3
- PsLD3: sum total alive and dead adult parasitoids at census t for Long-duration treatment, replicate microcosm 3
- HaLD5: number of adult weevils alive at census t for Long-duration treatment, replicate microcosm 5
- HdLD5: number of adult weevils dead at census t for Long-duration treatment, replicate microcosm 5
- HsLD5: sum total alive and dead adult weevils at census t for Long-duration treatment, replicate microcosm 5
- PaLD5: number of adult parasitoids alive at census t for Long-duration treatment, replicate microcosm 5
- PdLD5: number of adult parasitoids dead at census t for Long-duration treatment, replicate microcosm 5
- PsLD5: sum total alive and dead adult parasitoids at census t for Long-duration treatment, replicate microcosm 5
- HaSD1: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 1
- HdSD1: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 1
- HsSD1: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 1
- PaSD1: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 1
- PdSD1: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 1
- PsSD1: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 1
- HaSD2: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 2
- HdSD2: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 2
- HsSD2: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 2
- PaSD2: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 2
- PdSD2: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 2
- PsSD2: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 2
- HaSD4: number of adult weevils alive at census t for Short-duration treatment, replicate microcosm 4
- HdSD4: number of adult weevils dead at census t for Short-duration treatment, replicate microcosm 4
- HsSD4: sum total alive and dead adult weevils at census t for Short-duration treatment, replicate microcosm 4
- PaSD4: number of adult parasitoids alive at census t for Short-duration treatment, replicate microcosm 4
- PdSD4: number of adult parasitoids dead at census t for Short-duration treatment, replicate microcosm 4
- PsSD4: sum total alive and dead adult parasitoids at census t for Short-duration treatment, replicate microcosm 4
File: power-period_repeated_measures.csv
Description: This dataset is derived from the R code: Time series analysis. It reports for each period, the power of that period based on a wavelet analysis. Codes for each column of data are as follows:
Variables
- Species: Host or Ptoid (Parasitoid) Treatment: treatment, either Control, long-duration treatment (LD), or short-duration treatment (SD)
- Repl: replicate microcosm number
- P2: the spectrum power for period 2 oscillations
- P3: the spectrum power for period 3 oscillations
- P4: the spectrum power for period 4 oscillations
- P5: the spectrum power for period 5 oscillations
- P6: the spectrum power for period 6 oscillations
- P7: the spectrum power for period 7 oscillations
- P8: the spectrum power for period 8 oscillations
- P9: the spectrum power for period 9 oscillations
- P10: the spectrum power for period 10 oscillations
- P11: the spectrum power for period 11 oscillations
- P12: the spectrum power for period 12 oscillations
- P13: the spectrum power for period 13 oscillations
- P14: the spectrum power for period 14 oscillations
- P15: the spectrum power for period 15 oscillations
- P16: the spectrum power for period 16 oscillations
File: Time_series_analysis2.R
Description: Time series for each microcosm and temporal block were plotted, host and parasitoid means and SDs were computed, and wavelet analyses were conducted using the R code
File: treatment_mean-SD_log10.csv
Description: This dataset is derived from the Time series analysis. R code and contains means and SDs for host and parasitoid time series, after deleting transient dynamics (the first 25 census periods). Codes for each column of data are as follows:
Variables
- Mesocosm: mesocosm number
- Group: temporal block (1 or 2)
- Species: host or parasitoid (ptoid)
- TRTMNT: Control, Long-distance (LD) or Short-distance (SD) treatment
- Mean: log10 mean number of individuals for that species and microcosm
- SD: log10 standard deviation, the number of individuals for that species and microcosm
File: Control_simulation_model_ndnp2.R
Description: Simulation model for the control treatment with beans added daily (no pulse)
File: Control_simulation_model2.R
Description: Simulation model for the control treatment with a pulse of beans every 12 days
File: Host_fate.R
Description: R code for analyzing the fate of moth beans in the experimental microcosms
File: Host_feeding.R
Description: R code for the analysis of the experimental study on host feeding by Anisopteromalus calandrae
File: LD_simulation_model2.R
Description: Simulation model for the long-duration treatment in which there is a 12-day pulse of beans and more intraspecific competition among hosts
File: LD_simulation_model_ndnp2.R
Description: Simulation model for the long-duration treatment with beans added daily (no pulse) and more intraspecific competition among hosts
File: Power-period_repeated_measures.R
Description: Using data on the power for each period from a wavelet analysis (see time series analysis), we conducted a Repeated-measures ANOVA, followed by separate one-way ANOVAs for each time period. The analyses were conducted using this R code
Fil.e: SD_simulation_model2.R
Description: Simulation model for the short-duration treatment with a pulse of beans every 12 days.
File: SD_simulation_model_ndnp2.R
Description: Simulation model for the short-duration treatment with beans added daily (no pulse).
Code/software
Data for all analyses are provided as *.csv files. All data analyses and models were written using R version 4.3.1.
Changes after Mar 31, 2026:
There was a minor bug in the R code for the models that does not affect the analyses reported in the manuscript. It involves a calculation of the total number host fed each day. The code calculates the number host fed with a one day delay, instead of the current number.
