DOI: 10.5061/dryad.np5hqbzx4
title: Data from: Bet hedging is not sufficient to explain germination patterns of a winter annual plant

description: Bet hedging consists of life history strategies that buffer against
 environmental variability by trading off immediate and long-term fitness.
 Delayed germination in annual plants is a classic example of bet hedging
 and is often invoked to explain low germination fractions. We examined
 whether bet hedging explains low and variable germination fractions among
 20 populations of the winter annual plant Clarkia
 xantiana ssp. xantiana that experience substantial
 variation in reproductive success among years. Leveraging 15 years of
 demographic monitoring and 3 years of field germination experiments, we
 assessed the fitness consequences of seed banks and compared optimal
 germination fractions from a density-independent bet-hedging model to
 observed germination fractions. We did not find consistent evidence of bet
 hedging or the expected trade-off between arithmetic and geometric mean
 fitness, though delayed germination increased long-term fitness in 7 of 20
 populations. Optimal germination fractions were 2 to 5 times higher than
 observed germination fractions, and among-population variation in
 germination fractions was not correlated with risks across the life cycle.
 Our comprehensive test suggests that bet hedging is insufficient to
 explain the observed germination patterns. Understanding variation in
 germination strategies will likely require integrating bet hedging with
 complementary forces shaping the evolution of delayed germination.
description: A full description of the methods for data
 collection are provided in the manuscript associated with this Dryad
 submission. Briefly, we used field surveys and experiments to observe
 components of above- and below-ground demography for 20 populations across
 the range of <em>Clarkia
 xantiana</em> ssp. <em>xantiana</em>. To collect data on
 seedling survival, fruit production, and seed set, we used field surveys.
 In each population, these surveys included observations in both permanent
 plots as well as additional, haphazardly sampled plots arrayed across the
 population. To observe emergence of seedlings and seeds remaining intact
 in the soil seed bank, we conducted field experiments, which were
 complemented with lab experiments in order to assay viability of seeds.
 Brief details for each dataset are provided below, and the manuscript
 associated with these datasets describes the survey and experimental
 methods in further detail. We also estimated cumulative precipitation from
 February-June at each of the 20 populations, following methods described
 in a manuscript previously published on this study system (Eckhart et al.
 2011). The
 <em>README.md</em> file describes the contents of each file in
 this Dryad repository. Two metadata files provide additional information
 about the files in this Dryad submission: (1)
 <em>attributes.csv</em> is a comma-separated value file that
 lists and defines each variable in all the data files, and
 (2) <em>creators.csv</em> is a comma-separated value file that
 lists the co-authors on the manuscript associated with this Dryad
 submission. The metadata files were created using the R package dataspice
 (Boettiger et al. 2021).  countFruitsPerPlantAllPlants.csv Observations of total fruit equivalents per plant from field surveys in 2006-2012. Data come from up to 15 plants per permanent plot, plus plants found across the site. The counts are of “undamaged fruit equivalents” per plant, and were made by counting the number of undamaged fruits, and then counting the damaged fruits and estimating how many undamaged fruits these damaged fruits amounted to. variableName description site Site acronym countFruitNumberPerPlant   Count of fruits on a plant; the count combined undamaged and damaged fruits into a single value when fruits were surveyed in the field permanentPlot Binary variable indicating whether the plant on which fruits were counted was growing in one of 30 permanent plots at the site (1=true) or located in either an additional, haphazardly located plot or found by surveying the site (0=false) damage Binary variable indicating whether the fruit that was used to count seeds was undamaged (0=false) or damaged (1=true); all observations in this dataset have NA here because the count is of total fruit equivalents year Year in which observations were made countFruitsPerPlantFromPermanentPlots.csv Observations of total fruit equivalents per plant from field surveys in 2007-2012. Data come from up to 15 plants per permanent plot. The counts are of “undamaged fruit equivalents” per plant, and were made by counting the number of undamaged fruits, and then counting the damaged fruits and estimating how many undamaged fruits these damaged fruits amounted to. variableName description site Site acronym transect Transect number position Plot number plantNumber Variable indexing the plants on which fruits per plant was recorded; the index starts at 1 in each permanent plot in each year at each site countFruitsPerPlant   Count of fruits on a plant; the count combined undamaged and damaged fruits into a single value when fruits were surveyed in the field year Year in which observations were made countSeedPerFruit.csv Observations of seeds per fruit from seeds collected in the field in 2006-2020. From 2006-2020, data are for counts of seeds per undamaged fruit for up to roughly 30 fruits per site. From 2013-2020, the data also include the counts of seeds per damaged fruit. variableName   description site Site acronym year Year in which observations were made damaged Binary variable indicating whether seeds were counted in a fruit that was undamaged (0=false) or damaged (1=true) sdno Count of seeds countUndamagedDamagedFruitsPerPlantAllPlants.csv Observations of undamaged and damaged fruits per plant from field surveys in 2013-2020. Data come from up to 15 plants per permanent plot, plus plants found across the site. The counts are of “undamaged fruits” and "damaged fruits" on each plant. variableName description site Site acronym countUndamagedFruitNumberPerPlant   Count of undamaged fruits on a plant countDamagedFruitNumberPerPlant Count of damaged fruits on a plant permanentPlot Binary variable indicating whether the plant on which fruits were counted was growing in one of 30 permanent plots at the site (1=true, 0=false) year Year in which observations were made countUndamagedDamagedFruitsPerPlantFromPermanentPlots.csv Observations of undamaged and damaged fruits per plant from field surveys in 2013-2020. Data come from up to 15 plants per permanent plot. The counts are of “undamaged fruits” and "damaged fruits" on each plant. variableName description site Site acronym transect Transect number position Plot number plantNumber Variable indexing the plants on which fruits per plant was recorded; the index starts at 1 in each permanent plot in each year at each site countUndamagedFruitsPerPlant   Count of undamaged fruits on a plant year Year in which observations were made countDamagedFruitsPerPlant Count of damaged fruits on a plant seedBagsData.csv Observations of intact seeds and germinants from the seed bag burial experiment, conducted from October 2005-October 2008. The experiment consisted of 3 experimental rounds, starting in October 2005, 2006, and 2007. The data in this file includes counts of intact seeds and germinants. The data for the lab component of the seed bag burial experiment is found in viabilityData.csv. variableName   description site Site acronym transect Transect number position Plot number bagNo Bag number round Experimental round in which the bag was buried yearStart Year in which the experimental round started; bags were buried In October of this year age Number of years the bag had been buried when it was collected from the field in October yearData Year in which observations were made for this bag; the bag was dug up in January and October of this year seedlingJan Count of seedlings in the bag when it was dug up in January intactJan Count of intact, ungerminated seeds in the bag when it was dug up in January totalJan Sum of the count of seedlings and intact, ungerminated seeds in the bag when it was dug up in January intactOct Count of intact seeds in the bag when it was dug up in October seedlingFruitingPlantCountsPermanentPlots.csv Observations of seedlings and fruiting plants in permanent plots from 2006-2020. Data are counts of seedlings in permanent plots in January/February censuses, and counts of fruiting plants in permanent plots in June/July censuses. variableName description site Site acronym transect Transect number position Plot number year Year in which observations were made seedlingNumber   Count of seedlings in 0.5 m x 1 m permanent plot; NA if not recorded fruitplNumber Count of fruiting plants in 0.5 m x 1 m permanent plot; NA if not recorded siteAbioticData.csv Summary of abiotic variables associated with each study site in the long-term study of <em>Clarkia xantiana</em> ssp. <em>xantiana</em> demography. variableName   description site Site acronym siteName Full name for site easting Geographic position reported as eastward measured distance, reported for UTM zone 11, NAD 1927 (units of 100 meters) northing Geographic position reported as northward measured distance, reported for UTM zone 11, NAD 1927 (units of 100 meters) elevation Height above sea level (meters) area Size of the <em>Clarkia xantiana</em> ssp. <em>xantiana</em> population at the study site (hectares) surfaceRock Dominant soil parent material springPrecipitationData.csv Cumulative spring precipitation at each of the study sites and years in the long-term study of <em>Clarkia xantiana</em> ssp. <em>xantiana</em> demography. A full description of the methods used to estimate cumulative precipitation at the study sites was previously published (Eckhart et al. 2011). variableName   description site Site acronym year Year in which observations were made springPrecipitation Cumulative precipitation from February to June, estimated from daily rainfall measurements at weather stations in the field and interpolated to the location of study populations (units of millimeters)  viabilityData.csv Observations from lab germination assays and viability assays that are associated with the seed bag burial experiment. The lab experiments were conducted with seeds from seed bags that were recovered from the field in October 2006, 2007, and 2008. The data for the field component of the seed bag burial experiment is found in seedBagsData.csv. variableName   description site Site acronym bagNo Bag number round Experimental round in which the bag was buried age Number of years the bag had been buried when it was collected from the field in October block Block number for the Petri dishes used in germination trials germStart Count of seeds starting the germination trials germCount Count of seeds germinating over the course of the germination trials viabStart Count of seeds tested in the viability trials viabStain Count of seeds staining red in the viability trials <strong>Works cited</strong> Boettiger, C., Chamberlain, S., Fournier, A., Hondula, K., Krystalli, A., Mecum, B., Salmon, M., Webbink, K., and Woo, K. (2021). dataspice: Create Lightweight Schema.org Descriptions of Data. R package version 1.1.0. https://CRAN.R-project.org/package=dataspice Eckhart, V. M., M. A. Geber, W. F. Morris, E. S. Fabio, P. Tiffin, and D. A. Moeller. (2011). The Geography of Demography: Long-Term Demographic Studies and Species Distribution Models Reveal a Species Border Limited by Adaptation. The American Naturalist 178:S26–S43. https://doi.org/10.1086/661782 
description: Data collection is described in the manuscript associated with
 this dataset. Code associated with the analysis of the
 raw data provided in this Dryad submission is submitted on Zenodo and can
 be found at the following link: https://doi.org/10.5281/zenodo.7734839 The corresponding authors for this dataset are Monica Geber (mag9@cornell.edu), David Moeller (moeller@umn.edu), and Vincent Eckhart (eckhart@grinnell.edu).