Multiple signaling increases both prey response and diversity in a carnivorous pitcher plant
Data files
Dec 05, 2025 version files 734.79 KB
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ColNectVol.csv
66.90 KB
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colNectVolSetUp.csv
34.86 KB
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contrast.csv
3.90 KB
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gcms2024-05_normalized.csv
1.04 KB
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ModelPreyProfile.csv
34.71 KB
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Paint_RealPlant_Spectra.csv
548.52 KB
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PostHocVOC.csv
12.62 KB
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README.md
13.22 KB
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RealPlantPreyProfile.csv
19.02 KB
Abstract
Organisms across the tree of life use multiple signals such as color, movement, sound, and many types of chemical signals to convey seemingly the same information. These signals are potentially costly, yet the repeated evolution of multiple signaling across systems suggests an adaptive value. Using prey attraction by a carnivorous pitcher plant (Sarracenia purpurea), we tested two non-mutually exclusive hypotheses of how multiple signaling may be adaptive: 1) the receiver variability hypothesis that different signals reach different receivers, increasing the diversity of the receiver response and 2) the redundant signal hypothesis, that additional signals increase the probability of signal transmission to relevant receiver(s). We used artificial pitcher plant models with red coloration, nectar, and volatile organic compound signals to examine both the receiver variability and redundant signal hypotheses in a manipulative, full factorial field experiment in S. purpurea native habitat. Like pitcher plants, models captured visiting arthropods, which pitcher plants digest to cope with nutrient-poor bog soils. We found support for both the receiver variability and redundant signal hypotheses; both color and nectar additively increased prey diversity, as well as increased the capture of the bog specialist ant Myrmica lobifrons. Synthesis: Our results suggest that increasing both signal redundancy and diversity of respondents are substantial benefits that balance the costs of multiple signaling.
Description of data and files
Describes data from artificial pitcher plants (modeled off of Sarracenia purpurea) placed in 4 Michigan pitcher plant bogs in August 2023, as well as several follow up experiments conducted in the summer of 2024. Artificial pitcher plants contain a presence/absence full factorial of three putative prey attracting signals: red coloration, extrafloral nectar, and artificial VOC blends. Data on the spectral reflectence of real plants and painted models are also included in this dataset.
Files
Raw data files:
ColNectVol.csv -- Abundance of each prey morphospecies in each model pitcher plant at each site. Includes only models that captured prey (see below).
colNectVolSetUp.csv -- contains a unique row for each artificial pitcher plant placed at each bog site, regardless of prey capture outcome. Also lists field work dates.
contrast.csv -- The background moss color where each artificial pitcher plant was placed at the Upper Manistee Headwaters Preserve Site for analysis of the influence of background contrast on prey capture.
gcms2024-05_normalized.csv -- GCMS peaks and calcaluated raw and normalized emission rates of each compound in the artifical VOC blend. Model pitcher plants were set up in a sunny, well ventilated greenhouse in the same way they were set up in the bog sites; silicone septa were soaked in VOC blend in DCM, then dried for 1 hour before being hung inside artificial pitcher plants. Models were bagged and sampled for 30 minutes starting at the specified hour post set up.
ModelPreyProfile.csv -- Abundance of each prey morphospecies in each model pitcher plant used to compare real plants and model plants. Includes only models that captured prey.
RealPlantPreyProfile.csv -- Abundance of each prey morphospecies in each real pitcher plant used to compare real plants and model plants.
Paint_RealPlant_Spectra.csv -- spectral reflectance data from real plants and painted model pitcher plants.
PosHocVOC.csv -- Abundance of each prey morphospecies in each model pitcher plant used in the post hoc VOC experiment testing prey response to the high VOC blend, the biologically realistic blend, and a solvent (DCM) control.
Appendix A.zip -- microscope images of each morphospecies referenced in ColNectVol.csv; also includes "InsectIDNotes.docx" file of descriptions of each insect, and the sample where that morphospecies was first identified.
Code:
MultipleSignalingAnalysis.Rmd -- Code for data analysis and plots of main hypothesis testing, including only data from red and nectar treatments.
MultipleSignalingAnalysis.html -- Knitted version of MultipleSignalingAnalysis.Rmd file
MultipleSignalingAnalysis_w_VOCs.Rmd -- Code for data analysis and plots of main hypothesis testing, including all data (red, nectar, and VOC treatments)
MultipleSignalingAnalysis_w_VOCs.html -- Knitted version of MultipleSignalingAnalysis_w_VOCs.Rmd
ModelValidation.Rmd -- Code for data analysis and plots related to model validation, VOC lure and GCMS calculations, post hoc VOC experiments and spectra comparison
ModelValidation.html -- Knitted version of ModelValidation.Rmd
Glossary:
ColNectVol.csv
- trt -- the experimental treatment applied to each model; null is -nect - col - vol; the capital letters R N and V refer to the presence of each signal (Red veins, nectar, and VOCs, respectively)
- rep -- replicate, or the microsite array each model was placed within. Replicates range from 1-30 at each site.
- site -- the site where each model was placed; YSDL = Yankee Springs Deep Lake Yankee Springs Deep Lake boat launch (42.620, -85.457),
- YSLL = Yankee Springs Long Lake (42.627, -85.505)
- error -- 1 = some kind of error occured, including a tube lost or spilled during collection. Blank cells indicate no error
- P -- the total number of prey items collected from each model pitcher plant; includes "NP" column (see below); used as an internal check during lab processing of samples
- NP -- 'no prey'; 1 = no prey were found after examination under a microscope. Blank cells indicate prey was found
- ref -- 1 = storage tube was removed from others as a morphospecies identification reference; blank cells indicate storage tube in main storage box
- Unk2:spider -- number of items of that morphospecies, species, or larger taxonomic group found in the artificial pitcher plant. Blank cells indicate none of that taxon was found. See paper Supplement for morphospecies descriptions and images.
- sarc -- the number of Fletcherymia fletcheri larvae (Sarcophagid flies specialist on pitcher plants phytotelmata) found in the artificial pitcher plant
- notes -- notes
colNectVolSetUp.csv
- trt -- the experimental treatment applied to each model; null is -nect - col - vol; the capital letters R N and V refer to the presence of each signal (Red veins, nectar, and VOCs, respectively)
- rep -- replicate, or the microsite array each model was placed within. Replicates range from 1-30 at each site.
- site -- the site where each model was placed; YSDL = Yankee Springs Deep Lake Yankee Springs Deep Lake boat launch (42.620, -85.457),
- KBSB = Kellogg Biological Station Sherman Farm Marsh (42.355, -85.372), UMHP = Upper Manistee Headwaters Preserve (44.745, -85.005)
- and GRNA Grass River Natural Area (44.921, -85.242).
- dateOut -- the date (month/day/year) the model was placed in the bog
- dateIn -- the date (month/day/year) the model was retrieved from the bog
- spill -- 1 = tube spilled during collection. Blank cells indicate no error
- error -- 1 = some kind of error occured, including a tube lost. Blank cells indicate no error
- tube -- 1 = an alcohol storage tube was created during initial sample processing: possible prey was visible in the bottom of the model plant. 0 = no alcohol storage tube was created, model
- considered to have captured no prey. The rows of the ColNectVol.csv sheet correspond to those rows of this sheet with a 1 in the tube column.
- notes -- notes
contrast.csv
- trt -- the experimental treatment applied to each model; null is -nect - col - vol; the capital letters R N and V refer to the presence of each signal (Red veins, nectar, and VOCs, respectively)
- rep -- replicate, or the microsite array each model was placed within. Replicates range from 1-30 at each site.
- site -- the site where each model was placed; UMHP = Upper Manistee Headwaters Preserve (44.745, -85.005)
- sphCol -- the color of the Sphagnum moss where each artificial pitcher plant was placed; either 'red' or 'green'
gcms2024-05_normalized.csv
- sample -- the sample code; +0 and +3 refer to hours post drying time (1 hour after removal from soaking solution) when the sample was taken
- R1-5 refer to replicate model pitcher plants 1 - 5, which were set up in a sunny greenhouse. EB refers to empty bag samples taken as a reference / control.
- Missing replicates are due to failed sample elutions due to low elution volumes
- time -- the time in hours post drying time (1 hour after removal from soaking solution) when the sample was taken
- rep -- replicate model pitcher plants 1 - 5; Missing replicates are due to failed sample elutions due to low elution volumes
- eb -- sample was an empty bag samples taken as a reference / control
- ocimene:total -- peak area for each compound, and the sum of all peak areas for the sample
- ocimene_nghr:total_nghr -- the emission of each compound and the total emission for each sample, in nanograms per hour
- ocimene_nghr_norm:total_norm -- the emission of each compound and the total emission for each sample (ng per hour), normalized by subtracting the emission of each compound found in the empty bag at that time point
ModelPreyProfile.csv
- trt -- the experimental treatment applied to each model; null is -nect - col - vol; the capital letters R N and V refer to the presence of each signal (Red veins, nectar, and VOCs, respectively)
- rep -- replicate, or the microsite array each model was placed within. Replicates range from 1-30 at each site.
- site -- the site where each model was placed; YSDL = Yankee Springs Deep Lake Yankee Springs Deep Lake boat launch (42.620, -85.457),
- KBSB = Kellogg Biological Station Sherman Farm Marsh (42.355, -85.372), UMHP = Upper Manistee Headwaters Preserve (44.745, -85.005)
- and GRNA Grass River Natural Area (44.921, -85.242).
- spill -- 1 = tube was spilled during collection. NA indicates tube was not spilled
- tube -- 1 = an alcohol storage tube was created during initial sample processing: possible prey was visible in the bottom of the model plant. 0 = no alcohol storage tube was created, model
- considered to have captured no prey.
- NP -- 'no prey'; 1 = no prey were found after examination under a microscope. NAs indicate prey was found.
- ref -- 1 = storage tube was removed from others as a morphospecies identification reference; blank cells indicate storage tube in main storage box
- unk3:ukn -- number of items of that morphospecies, species, or larger taxonomic group found in the artificial pitcher plant. Blank cells indicate none of that taxon was found. See paper Supplement for morphospecies descriptions and images.
- notes -- notes
RealPlantsPreyProfile.csv
- plantID -- plant identifier; number is unique within a site, no correspondence between plants with the same ID at different sites. Number corresponds to replicate number of closest artificial pitcher plant replicate
- initial_final -- i = initial, or prey collected when models placed in bog on day one of data collection (corresponds to diversity of prey capture over the previous weeks and months); f = final, prey collected from plant when model pitcher plants
- were retrieved 24 hours later (corresponds to prey capture by real plants over the same time interval as the model pitcher plants)
- site -- the site where each model was placed; YSDL = Yankee Springs Deep Lake Yankee Springs Deep Lake boat launch (42.620, -85.457),
- KBSB = Kellogg Biological Station Sherman Farm Marsh (42.355, -85.372), UMHP = Upper Manistee Headwaters Preserve (44.745, -85.005)
- and GRNA Grass River Natural Area (44.921, -85.242).
- spill -- 1 = tube was spilled during collection. NA indicates tube was not spilled
- NP -- 'no prey'; 1 = no prey were found after examination under a microscope. NAs indicate prey was found.
- globSpring:unk -- number of items of that morphospecies, species, or larger taxonomic group found in the artificial pitcher plant. Blank cells indicate none of that taxon was found. See paper Supplement for morphospecies descriptions and images.
PostHocVOC.csv
- trt -- the experimental treatment applied to each model; null is -nect - col - vol; the capital letters R N and V refer to the presence of each signal (Red veins, nectar, and VOCs, respectively)
- rep -- replicate, or the microsite array each model was placed within. Replicates range from 1-30 at each site.
- tube -- 1 = an alcohol storage tube was created during initial sample processing: possible prey was visible in the bottom of the model plant. 0 = no alcohol storage tube was created, model
- considered to have captured no prey.
- finished -- internal check that tube was processed (sum of notube, NP, and prey taxon columns)
- notube -- 1 = no alcohol storage tube was created; NA indicates a storage tube was created
- NP -- no prey, 1 = after examination under a microscope, no prey items were found; NA indicates prey was found, or no alcohol storage tube was created
- unknown:snail -- abundance of each category of prey
- notes -- notes
- sum -- total count of all prey captured
- bin -- binary prey capture; 1 = some kind of prey captured, 0 = no prey was captured
Paint_RealPlant_Spectra.csv -- spectral reflectance data from real plants and painted model pitcher plants
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wl -- the sampling wavelength, in nm
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redPaint -- samples of red areas of painted models ('Burgundy', Plaid acrylic paint); columns include three samples (1-3) and three measurements per sample
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greenPaint -- samples of green areas of painted models ('Fresh Foliage', Plaid acrylic paint); columns include three samples (1-3) and three measurements per sample
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All other columns named with the following conventions:
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rd010, rd014, and rd002 all refer to individual pitcher plants growing in the Gilbert Lab green house living collection
- 'vase' refers to the tubular body of the pitcher (colored red in all sampled plants)
- 'grSpot' refers to a green area of the inner hood of the pitcher leaf
- 'rdSpot' refers to a red area of the inner hood of the pitcher leaf
- 'young' modifies some samples to indicate the leaf was young and not fully pigmented or lignified
Each sample was measured three times, with the columns bearing identical names. Suffixes "", ".1", and ".2" differentiate between three replicate measurements for each sample.
Required packages:
tidyverse, glmmTMB, performance, DHARMa, brms, tidybayes, vegan, pavo, viridis, ggpubr
