Data from: Food-plant-associated predation risk and its impact on larval development and adult morphology in a specialist butterfly
Data files
Apr 08, 2026 version files 1.04 MB
-
2018_VR_Predation_CG_subsetAfAs.csv
133.74 KB
-
2018_VR_Predation_CG_SurvCurve.csv
9.35 KB
-
2019_VR_Predation_CG_subsetAfAs.csv
49.06 KB
-
2019_VR_Predation_CG_SurvCurve.csv
5 KB
-
Dryad_EcoEnto_2026.R
60.05 KB
-
fulldays_kestrel_DA.csv
578.11 KB
-
kestrel_RN.csv
176.12 KB
-
README.md
31.64 KB
Abstract
Predation influences herbivore behavior, physiology, and reproductive success through both direct consumption and non-lethal effects arising from predation risk alone. Plant traits, including defensive traits and architecture, can also influence herbivores both directly and indirectly, either by affecting feeding efficiency or by affecting herbivores' risk of predation. Yet, it remains unclear how predators and plant traits interact to shape the developmental outcomes of larval herbivores, particularly under field conditions. To understand larval performance within a tritrophic framework, we investigated how predation risk and food-plant identity influence caterpillar development and subsequent adult morphology using the monarch butterfly (Danaus plexippus) as a focal species. We compared responses to predation risk in monarch caterpillars feeding on two common species of western milkweed, Asclepias speciosa and Asclepias fascicularis, which differ in various traits, including defensive traits and architectural complexity. Using an outdoor common garden experiment in two growing seasons, caterpillars were assigned to plants of each of the two species with or without a predator-exclusion cage. Interestingly, we observed trends of slower weight gain in caterpillars feeding on A. speciosa than in those feeding on A. fascicularis, but only when caterpillars were exposed to predators. Additionally, in the first year of our study, predator exposure increased caterpillar development time on A. speciosa, but not on A. fascicularis. Predator exposure on A. speciosa was associated with reduced larval weight gain in the second year of our study, and because larval weight gain strongly predicted adult wing area, this pattern suggests a potential indirect effect of predators on wing size. In contrast, predator exposure had no detectable effect on wing morphometrics of adults that had developed on A. fascicularis. Our findings highlight that larval food-plant species can modulate the magnitude of predator effects at multiple life stages of an insect herbivore. This demonstrates that predator-exposure induced developmental changes can depend on the identity of the larval food-plant and can carry forward to affect adult morphology, with potential implications for the fitness of migratory insects.
Dataset DOI: 10.5061/dryad.51c59zwgx
Description of the data and file structure
These data were collected from two years (2018–2019) of common-garden experiments assessing the effects of larval host-plant species and predator exposure on monarch butterfly (Danaus plexippus) development and adult morphology. Monarch larvae were tracked daily from second instar until adulthood or death on either predator-exclusion or control plants, with one larva per plant. Larval survival, developmental timing, and pre-pupal weight gain were recorded. Adults that emerged were checked for OE infection, weighed, and measured for wing morphometrics, including length, width, area, loading, and roundness. Plant architecture was also recorded prior to larval introduction.
Files and variables
File: 2019_VR_Predation_CG_subsetAfAs.csv
Description: Data from second year of a two-year common-garden experiment tracking monarch (Danaus plexippus) larvae on two milkweed species (Asclepias fascicularis, A. speciosa) under predator-exposure treatments. Larvae were monitored daily from second instar to adult emergence or death. Data is summarized into columns for developmental timing, morphology, parasite infection, and with details on predators present as well as relevant plant traits. The dataset is structured to support analyses of larval performance, predator effects, and plant interactions within a tritrophic framework
Variables
- plant_location: Location of the plant in the common garden, labeled as row (R##) and plant within row (P##). Please note that numbers are not always consecutive due to some plants not surviving. Categorical
- monarch_id: Categorical unique identifier for each caterpillar, including year of experiment and plant location.
- milkweed_spp: Species of milkweed (Af = Asclepias fascicularis and As = A. speciosa) grown from seeds collected from multiple wild populations in northern Nevada and California. Categorical.
- treatment: Experimental treatment-level, either "roof" (open-air, predator-exposed) or "cage" (predator-exclusion). Categorical.
- cat_into_field: Date the caterpillar was placed onto the assigned plant in the common garden (MM/DD/YYYY).
- larva_last_day_seen: Last day the caterpillar was observed on its assigned plant (or date last seen as larva before pupation) in MM/DD/YYYY format.
- larval_time_in_field: Number of days the caterpillar was alive in the garden (numeric). If missing immediately after placement, counted as 0.5 days. Numeric.
- death_type_roof: Type of death as observed for caterpillars in the “roof” (open control) treatment. Possible values:
missing,chewed,black,sucked,shriveled,parasitism.NAif not applicable, if caterpillar survived. Categorical. - death_type_cage: Type of death as observed caterpillars in the predator-exclusion "cage" treatment. Possible values:
shriveled,black,failed_pupa.NAif not applicable, if caterpillar survived. Categorical. - instar_at_death: Last larval stage observed (2–5 or pupa).
NAif individual survived to adulthood. Categorical. - egg_date: Estimated oviposition date of the egg for plants in greenhouse in MM/DD/YYYY format.
- x2_instar_g: Weight in grams of the caterpillar at second instar prior to placement into experimental garden. Numeric.
- x2_inst_mm_avg: Average of three length measurements (mm) at time of weighing for second instar. Numeric.
- date_3rd_instar: Date the caterpillar was first observed as new to early third instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - date_4th_instar: Date the caterpillar was first observed as new to early fourth instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - days_3to4: Number of days between third and fourth instars.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - garden_days_to_4: Number of days from placement into the common garden to reaching early fourth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4_inst_g: Weight of caterpillar at fourth instar (g).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_g/d: Fourth instar growth rate in grams per day, calculated as
(final mass at the start of the fourth instar minus the larva’s mass measured at second instar prior to placement) divided by the number of days the larva spent in the garden up to the fourth instar. Numeric. - x4th_instar_length1_mm: First length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_instar_length2_mm: Second length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_instar_length3_mm: Third length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed or if the caterpillar could not be measured a third time. Numeric. - x4th_inst_avg_length: Average of length measurements for fourth instar (mm).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - caged_4: Y/N if caterpillar was caged in at fourth instar to prevent movement between plants. Categorical.
- x5th_inst_date: Date the caterpillar was first observed as new to early fifth instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - days_4_to_5: Number of days between fourth and fifth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - days_garden_to_5th: Number of days from placement into the common garden to reaching early fifth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x5_inst_g: Weight of caterpillar at new or early fifth instar (g).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x5th_g/d: Fifth instar growth rate in grams per day, calculated as
(final mass at the start of the fourth instar minus the larva’s mass measured at second instar prior to placement) divided by the number of days the larva spent in the garden up to the fourth instar. Numeric. - x5th_instar_avg_length: Average of length measurements for fourth instar (mm).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - date_last_weighed: Date the caterpillar was last weighed as a late fifth instar that had ceased eating (pre-pupal), in DD/MM/YYYY format.
NAif the caterpillar did not reach this stage or if no final weight was obtained. - last_cat_weight_g: The larva’s final mass recorded in the garden before pupation or death, measured after feeding cessation but prior to pupal formation. This value represents the pre-pupal body mass used in calculating final growth rate.
NAif caterpillar did not survive to this stage or if a last weight was not able to be taken. Numeric. - final_rate_larval_weight_gain: Larval growth rate for time in common garden in grams per day. Calculated as the pre-pupal mass (after feeding cessation and gut voiding) minus the larva’s 2nd-instar mass at placement into the garden, divided by the number of days between those two measurements. Numeric.
- larval_surv_binary: 1 if caterpillar survived to be pre-pupal, 0 if not. Binary.
- date_pupated: Date larva began pupation in DD/MM/YYYY format.
- pupated_binary: 1 if late 5th instar pupated successfully, 0 if not.
NAif larva died before late 5th instar. Binary. - garden_days_to_pupa: Number of days from placement in the common garden to pupation.
NAif caterpillar did not survive to pupation. Numeric. - pupae_weight_g: Weight of pupa (g) after 24-48 h to allow hardening.
NAif caterpillar did not pupate. Numeric. - emerged_if_pupated: 1 if a pupa successfully eclosed as an adult, 0 if not.
NAif caterpillar did not pupate. Binary. - eclosed_date: Date of adult emergence in DD/MM/YYYY format.
NAif caterpillar did not survive. - days_garden_to_adult: Days from placement in garden (2nd instar) to adult eclosion.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - survived_binary: 1 if the caterpillar survived from placement into the common garden through to adult emergence, 0 if it died at any point before emergence. Binary.
- adult_sex: Male or female, if adult emerged.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Categorical. - adult_wetweight_g: Adult weight in grams from living butterfly after emergence and wing drying.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - adult_dryweight_g: Adult weight in grams from freeze-dried butterfly.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - adult_weight_diff: Difference between wet and dry weight in grams.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - drywt_gain_g/d: Adult dry weight gain per day (g/day), standardized by developmental period.
NAif caterpillar did not survive. Numeric. - adult_OEscore: Level of Ophryocystis elektroscirrha (OE) infection scored 0–5 based on spore density. Numeric.
- adult_Oebinary: 1 if any OE spores detected, 0 if none. Binary.
- wing_length_mm: Forewing length (mm) measured from removed and scanned wing using ImageJ. Numeric.
- wing_width_mm: Forewing width (mm) measured from removed and scanned wing using ImageJ. Numeric.
- wing_area: Forewing area (mm2) measured from removed and scanned wing using ImageJ. Numeric.
- perimeter_mm: Forewing perimeter (mm) measured from removed and scanned wing using ImageJ. Numeric.
- aspect_ratio: Forewing length divided by width. Numeric.
- roundness: 4 × [area / (π × length)^2^]; numeric measure of wing shape.
- wing_loading: Dry body mass (g) / wing area (mm2); indicates flight performance. Numeric.
- caged_cat_died: Y/N; if caterpillar in cage treatment died. Categorical.
- larvae_sick: Y/N; if caterpillar appeared sickly. Categorical.
- parasitized_pupa: Y if tachinid fly pupa emerged from monarch pupa; N if not.
NAfor cage treatment or non-pupated caterpillar. Categorical. - parastized_binary: 1 if tachinid fly pupa emerged from monarch pupa; 0 if not.
NAfor cage treatment or non-pupated caterpillar. Binary. - tachinid_surv_binary: 1 if tachinid pupa successfully produced an adult fly, 0 if not. Binary.
- predator_present_binary: 1 if any known-monarch predator was observed on the plant during the daily standardized check (wasp, ant, spider, mantis, milkweed bug, etc.), 0 if none were observed. Binary.
- predator_type1: Common name of first predator observed. Categorical.
- predator_type2: Common name of second predators observed. Categorical.
- predator_count: Number of predators observed on the plant. Numeric.
- aphids_binary: 1 if aphids present, 0 if not.
NAif in cage treatment. Binary. - plant_ID: Lab-specific code for plant lineage (seed collection site – mother – father – species). Categorical.
- plant_natal_code: Seed collection site abbreviation (e.g., BM = Battle Mountain, CA = California mix, RN = Reno). Categorical.
- plant_size: Size of plant at placement of larva in the garden. Categorical.
- plant_morphology: Details of plant morphology (e.g., bushy, spindly, or diseased). Categorical.
- branch_num_all: Total number of stems >2 inches on plant before the start of the experiment for all plants in the experiment. Numeric.
- branch_num_sub: Number of stems >2 inches at the start before the experiment for roof treatment only;
NAfor cages. Numeric. - cat_moved: Y/N if larva was moved from original plant to plant of same species due to plant die back. Categorical.
- plant_fungus: 1 if the plant showed signs of fungal infection, 0 if not. Binary.
- plant_flower: 1 if the plant was flowering, 0 if not; recorded only for roof treatment plants,
NAfor cage treatment. Binary.
File: 2018_VR_Predation_CG_subsetAfAs.csv
Description: Data from first year of a two-year common-garden experiment tracking monarch (Danaus plexippus) larvae on two milkweed species (Asclepias fascicularis, A. speciosa) under predator-exposure treatments. Larvae were monitored daily from second instar to adult emergence or death. Data is summarized into columns for developmental timing, morphology, parasite infection, and with details on predators present as well as relevant plant traits. The dataset is structured to support analyses of larval performance, predator effects, and plant interactions within a tritrophic framework
Variables
- plant_location: Location of the plant in the common garden, labeled as row (R##) and plant within row (P##). Please note that numbers are not always consecutive due to some plants not surviving. Categorical.
- monarch_id: Unique identifier for each caterpillar, including year of experiment and plant location. Categorical.
- milkweed_spp: Species of milkweed (Af= Asclepias fascicularis and As = A. speciosa) grown from seeds collected from multiple wild populations in northern Nevada and California. Categorical.
- treatment: Experimental treatment-level, either "roof" (open-air, predator-exposed) or "cage" (predator-exclusion). Categorical.
- cat_into_field: Date the caterpillar was placed onto the assigned plant in the common garden (MM/DD/YYYY).
- 30-Sep–8-Nov: Columns that reflect the daily observations of each caterpillar, coded by status such as out (day of placement into garden), miss (not found in standardized search time), surv (alive), caged (checked enclosure through mesh but larva not seen; cage not opened), presumed_dead (missing for consecutive days and considered predated), or
NAif not checked that day due to weather or other constraints. Categorical. - larva_last_day_seen: Last day the caterpillar was observed on its assigned plant (or date last seen as larva before pupation) in MM/DD/YYYY format.
- larval_time_in_field: Number of days the caterpillar was alive in the garden (numeric). If missing immediately after placement, counted as 0.5 days. Numeric.
- death_type_roof: Type of death as observed for caterpillars in the “roof” (open control) treatment. Possible values: missing, chewed, black, sucked, shriveled, parasitism.
NAif not applicable, if caterpillar survived. Categorical. - death_type_cage: Type of death as observed caterpillars in the predator-exclusion "cage" treatment. Possible values: shriveled, black, failed_pupa.
NAif not applicable, if caterpillar survived. Categorical. - instar_at_death: Last larval stage observed (2–5 or pupa).
NAif individual survived to adulthood. Categorical. - egg_date: Estimated oviposition date of the egg in MM/DD/YYYY format.
- x2_instar_g: Weight in grams of the caterpillar at second instar prior to placement into experimental garden. Numeric.
- x2_inst_mm_avg: Average of three length measurements (mm) at time of weighing for second instar. Numeric.
- date_3rd_instar: Date the caterpillar was first observed as new to early third instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - date_4th_instar: Date the caterpillar was first observed as new to early fourth instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - days_3to4: Number of days between third and fourth instars.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - garden_days_to_4: Number of days from placement into the common garden to reaching early fourth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4_inst_g: Weight of caterpillar at fourth instar (g).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_g/d: Fourth instar growth rate in grams per day, calculated as
(final mass at the start of the fourth instar minus the larva’s mass measured at second instar prior to placement) divided by the number of days the larva spent in the garden up to the fourth instar. Numeric. - x4th_instar_length1_mm: First length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_instar_length2_mm: Second length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x4th_instar_length3_mm: Third length measurements for fourth instar (mm) if weighed.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed or caterpillar could not measured a third time. Numeric. - x4th_inst_avg_length: Weight of caterpillar at fourth instar (g).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - caged_4: Y/N if caterpillar was caged in at fourth instar to prevent movement between plants. Categorical.
- x5th_inst_date: Date the caterpillar was first observed as new to early fifth instar in MM/DD/YYYY format.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. - days_4_to_5: Number of days between fourth and fifth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - days_garden_to_5th: Number of days from placement into the common garden to reaching early fifth instar.
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x5_inst_g: Weight of caterpillar at new or early fifth instar (g).
NAif caterpillar did not survive to this instar or if the beginning of this stage was not observed. Numeric. - x5th_g/d: Fifth instar growth rate in grams per day, calculated as
(final mass at the start of the fourth instar minus the larva’s mass measured at second instar prior to placement) divided by the number of days the larva spent in the garden up to the fourth instar. Numeric. - date_last_weighed: Date the caterpillar was last weighed as a late fifth instar that had ceased eating (pre-pupal), in DD/MM/YYYY format.
NAif the caterpillar did not reach this stage or if no final weight was obtained - last_cat_weight_g: The larva’s final mass recorded in the garden before pupation or death, measured after feeding cessation but prior to pupal formation. This value represents the pre-pupal body mass used in calculating final growth rate.
NAif caterpillar did not survive to this stage or if a last weight was not able to be taken. Numeric. - final_rate_larval_weight_gain: Larval growth rate for time in common garden in grams per day. Calculated as the pre-pupal mass (after feeding cessation and gut voiding) minus the larva’s 2nd-instar mass at placement into the garden, divided by the number of days between those two measurements. Numeric.
- larval_surv_binary: 1 if caterpillar survived to be pre-pupal, 0 if not. Binary.
- date_pupated: Date larva began pupation in DD/MM/YYYY format.
- pupated_binary: 1 if late 5th instar pupated successfully, 0 if not.
NAif larva died before late 5th instar. Binary. - garden_days_to_pupa: Number of days from placement in the common garden to pupation.
NAif caterpillar did not survive to pupation. Numeric. - pupae_weight_g: Weight of pupa (g) after 24-48 h to allow hardening.
NAif caterpillar did not pupate. Numeric. - emerged_if_pupated: 1 if a pupa successfully eclosed as an adult, 0 if not.
NAif caterpillar did not pupate. Binary. - eclosed_date: Date of adult emergence in DD/MM/YYYY format.
NAif caterpillar did not survive. - days_garden_to_adult: Days from placement in garden (2nd instar) to adult eclosion.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - survived_binary: 1 if the caterpillar survived from placement into the common garden through to adult emergence, 0 if it died at any point before emergence. Binary.
- adult_sex: Male or female, if adult emerged.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Categorical. - adult_wetweight_g: Adult weight in grams from living butterfly after emergence and wing drying.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - adult_dryweight_g: Adult weight in grams from freeze-dried butterfly.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - adult_weight_diff: Difference between wet and dry weight in grams.
NAif caterpillar did not successfully eclose or died prior to reaching pupation. Numeric. - drywt_gain_g: Adult dry weight gain per day (g/day), standardized by developmental period.
NAif caterpillar did not survive. Numeric. - adult_OEscore: Level of Ophryocystis elektroscirrha (OE) infection scored 0–5 based on spore density. Numeric.
- adult_Oebinary: 1 if any OE spores detected, 0 if none. Binary.
- wing_length_mm: Forewing length (mm) measured from removed and scanned wing using ImageJ. Numeric.
- wing_width_mm: Forewing width (mm) measured from removed and scanned wing using ImageJ. Numeric.
- wing_area: Forewing area (mm2) measured from removed and scanned wing using ImageJ. Numeric.
- perimeter_mm: Forewing perimeter (mm) measured from removed and scanned wing using ImageJ. Numeric.
- aspect_ratio: Forewing length divided by width. Numeric
- roundness: 4 × [area / (π × length)^2^]; numeric measure of wing shape.
- wing_loading: Dry body mass (g) / wing area (mm2); indicates flight performance. Numeric.
- plant_ID: Lab-specific code for plant lineage (seed collection site – mother – father – species). Categorical.
- plant_natal_code: Seed collection site two-letter abbreviation (e.g., BM = Battle Mountain, CA = California mix, RN = Reno). Categorical.
- predator_present_binary: 1 if any known-monarch predator was observed on the plant during the daily standardized check (e.g., wasp, ant, spider, mantis, milkweed bug, etc.), 0 if none were observed. Binary.
- predator_type1: Common name of first predator observed. Categorical.
- predator_type2: Common name of second predators observed. Categorical.
- predator_count: Number of predators observed on the plant. Numeric.
- aphids_binary: 1 if aphids present, 0 if not.
NAif in cage treatment. Binary. - larvae_sick: Y/N; if caterpillar appeared sickly. Categorical.
- parasitized_pupa: Y if tachinid fly pupa emerged from monarch pupa; N if not.
NAfor cage treatment or non-pupated caterpillars. Categorical. - parastized_binary: 1 if tachinid fly pupa emerged from monarch pupa; 0 if not.
NAfor cage treatment or non-pupated caterpillars. Binary. - tachinid_surv_binary: 1 if tachinid pupa successfully produced an adult fly, 0 if not.
NAfor non-parastized caterpillars. Binary. - plant_fungus: 1 if the plant showed signs of fungal infection, 0 if not. Binary.
- plant_flower: 1 if the plant was flowering, 0 if not; recorded only for roof treatment plants,
NAfor cage treatment. Binary. - plant_morphology: Details of plant morphology (e.g., bushy, spindly, or diseased). Categorical.
- notes: Relevant notes. Categorical.
File: kestrel_RN.csv
Description: Hourly microclimate measurements collected using Kestrel DROP D2 wireless data loggers placed inside predator-exclusion cages and mock-cage controls during follow-up field trials in Reno, NV. These measurements were used to quantify treatment-level differences in temperature and humidity within the caging structures.
Variables
- Device_Name: Identifier assigned to the Kestrel device. Categorical.
- Date: Calendar date of measurement (MM/DD/YYYY).
- FORMATTED.DATE_TIME: Timestamp in device-export format. Datetime.
- Time_Military: Time of day in 24 hr format (HH:MM). Time.
- Temperature_F: Air temperature in degrees Fahrenheit calculated by the device. Numeric.
- Relative_Humidity_Percent: Relative humidity (%) calculated by the device. Numeric.
- Heat_Index_F: Heat index (F) calculated by the device. Numeric.
- Dew_Point_F: Dew point (F) calculated by the device. Numeric.
- Duration_Min: Logger runtime at that record (min). Numeric.
- Treatment_Type: Experimental treatment-level, either "roof" (open-air, predator-exposed) or "cage" (predator-exclusion). Categorical.
- Device_Model: Model designation from the Kestrel record (DROP D2). Categorical.
- Data.Type: Indicates the record class in the Kestrel export, where “point” denotes a direct instantaneous sensor measurement taken at the logged interval.. Categorical.
- Date_Format_yyyy.MM.dd.hh.mm.ss.a: Timestamp in alternative device-export formatting. Datetime.
File: fulldays_kestrel_DA.csv
Description: Hourly microclimate measurements collected using Kestrel DROP D2 wireless data loggers placed inside predator-exclusion cages and mock-cage controls during follow-up field trials in Davis, CA. These measurements were used to quantify treatment-level differences in temperature and humidity within the caging structures.
Variables
- Device_Name: Identifier assigned to the Kestrel device. Categorical.
- Date: Calendar date of measurement (MM/DD/YYYY).
- FORMATTED.DATE_TIME: Timestamp in device-export format. Datetime.
- Time_Military: Time of day in 24 hr format (HH:MM). Time.
- Temperature_F: Air temperature in degrees Fahrenheit calculated by the device. Numeric.
- Relative_Humidity_Percent: Relative humidity (%) calculated by the device. Numeric.
- Heat_Index_F: Heat index (F) calculated by the device. Numeric.
- Dew_Point_F: Dew point (F) calculated by the device. Numeric.
- Duration_Min: : Logger runtime at that record (min). Numeric.
- Treatment_Type: Experimental treatment-level, either "roof" (open-air, predator-exposed) or "cage" (predator-exclusion). Categorical.
- Milkweed_sp: Milkweed species where the logger was placed (Af = A. fascicularis, As = A. speciosa). Categorical.
- Device_Model: Model designation from the Kestrel record (DROP D2). Categorical.
- Data.Type: Indicates the record class in the Kestrel export. “point” denotes a direct instantaneous sensor measurement taken at the logged interval.
- Date_Format_yyyy.MM.dd.hh.mm.ss.a: Timestamp in alternative device-export formatting. Datetime.
- Temperature_C: Temperature converted to Celsius. Numeric.
- DateTime: Combined standardized datetime.
File: 2018_VR_Predation_CG_SurvCurve.csv
Description: Dataset formatted for survival analysis of monarch caterpillars from the first year of a two-year common-garden experiment. The dataset summarizes time-to-event outcomes for larvae exposed to different predation-risk treatments on two milkweed species. These data were used to generate Kaplan–Meier survival curves and associated estimates.
Variables
- location: Unique identifier for caterpillar based on plant location in the common garden, formatted as R##P##. Categorical.
- milkweed_spp: Milkweed species that caterpillar was assigned to (Af = A. fascicularis, As = A. speciosa).
- treatment: Experimental treatment-level, either "R" for open-air, predator-exposed ("Roof") or "C" for predator-exclusion ("Cage"). Categorical.
- treatment_split: Categorical variable combining milkweed species and predation-risk treatment. Codes follow the pattern [species]_[predation], where Af = Asclepias fascicularis, As = A. speciosa, preds = predator-exposure (mock cage), and no_preds = predator-exclusion cage.
- time1: Time to event in days from placement until death or censoring for survival.
- event1: Event indicator (1 = death, 0 = censored). Binary.
- survived_binary: Survival outcome (1 = survived to adult, 0 = did not survive). Binary.
File: 2019_VR_Predation_CG_SurvCurve.csv
Description: Dataset formatted for survival analysis of monarch caterpillars from the second year of a two-year common-garden experiment. The dataset summarizes time-to-event outcomes for larvae exposed to different predation-risk treatments on two milkweed species. These data were used to generate Kaplan–Meier survival curves and associated estimates.
Variables
- location: Unique identifier for caterpillar based on plant location in the common garden, formatted as R##P##. Categorical.
- milkweed_spp: Milkweed species that caterpillar was assigned to (Af = A. fascicularis, As = A. speciosa).
- treatment: Experimental treatment-level, either "R" for open-air, predator-exposed ("Roof") or "C" for predator-exclusion ("Cage"). Categorical.
- treatment_split: Categorical variable combining milkweed species and predation-risk treatment. Codes follow the pattern [species]_[predation], where Af = Asclepias fascicularis, As = A. speciosa, preds = predator-exposure (mock cage), and no_preds = predator-exclusion cage.
- time1: Time to event in days from placement until death or censoring for survival.
- event1: Event indicator (1 = death, 0 = censored). Binary.
- survived_binary: Survival outcome (1 = survived to adult, 0 = did not survive). Binary
File: Dryad_EcoEnto_2026.R
Description: R script to clean and process monarch caterpillar datasets from 2018 and 2019 common garden experiments, calculate larval growth and survival metrics, data analyses and produce figures (including to generate Kaplan–Meier survival curves). The script also incorporates post-hoc Kestrel data logger measurements of temperature and relative humidity for microclimate assessment within predator-exclusion and control cages. This script uses all CSV files provided in the submission.
Code/software
All datasets can be viewed with any standard CSV reader, including open-source options. All analyses associated with these datasets were conducted in R (R Core Team) with RStudio. The following packages were used in the workflow: tidyverse, ggplot2, ggpubr, sjPlot, ggforce, car, MuMIn, DHARMa, survival, survminer, lmtest, lavaan, lavaanPlot, effects, lme4, lubridate, emmeans, and boot. The analysis workflow consisted of reading CSV files into R, performing data cleaning and transformations, fitting mixed-effects models, survival models, and structural equation models, evaluating model diagnostics, and generating figures.
Access information
Other publicly accessible locations of the data:
- None. The data have not been deposited elsewhere.
Data was derived from the following sources:
- All data were generated by the authors during the 2018 and 2019 common garden experiments and subsequent Kestrel microclimate trials. No external datasets were used.
We conducted a 2x2 factorial common garden experiment over two consecutive years (2018, N = 322; 2019, N = 171) in Reno, NV, to assess the effects of predation risk and host-plant identity (Asclepias speciosa and A. fascicularis) on monarch butterfly (Danaus plexippus) larval development and adult morphology. Predation risk was manipulated using predator-exclusion cages and open-sided "mock-cages" (referred to as "roof" in the dataset). Gardens were planted each year with first-year milkweed plants grown from seeds collected from multiple wild populations to account for intraspecific variation in plant chemistry and phenotype. Monarch larvae (F2 generation from local wild-caught females) were placed individually on plants and tracked daily from the second instar until adulthood or death. We recorded larval survival, developmental stage progression, and the timing of reaching the fifth instar. Adults were measured for dry weight and wing morphology (length, width, area, and roundness) using ImageJ. To monitor microclimatic differences between treatments, Kestrel DROP D2 wireless data loggers recorded hourly temperature and relative humidity at two locations (Reno and Davis). Larvae that suffered mortality from accidental causes (n = 2) were omitted from survival analyses but retained in developmental analyses.
Data were manually recorded in field notebooks, entered into spreadsheets, and verified for accuracy. All data manipulation and statistical analyses were conducted in R and RStudio using tidyverse, ggplot2, ggpubr, sjPlot, ggforce, glmmTMB, car, MuMIn, DHARMa, survival, survminer, lmtest, lavaan, lavaanPlot, effects, lme4, lubridate, emmeans, and boot. Statistical modeling included generalized linear models (binomial for survival), linear mixed-effects models (for microclimate, accounting for device ID as a random effect), and structural equation modeling (SEM).
