DOI: 10.6078/d1kx30
title: Data from: Topographic heterogeneity lengthens the duration of pollinator resources

description: The availability of sufficient and diverse resources across time is
 important for maintenance of biodiversity and ecosystem functioning. In
 this study we examine the potential for variation in environmental
 conditions across topographic gradients to extend floral resource timing.
 Flowering time on a landscape may vary across topography due to
 differences in abiotic factors, species turnover, or genotypic
 differences. However, the extent to which this variation in phenology
 affects overall flowering duration on a landscape, and the components of
 diversity that influence flowering duration, are unexplored. We
 investigate whether differences in flowering time due to topography yield
 an overall extension in duration of flowering resources in a northern
 California grassland. We recorded flowering time of pollinator resource
 species across four successive spring growing seasons (2015-2018) on
 paired north and south aspects. Flowering time differences were evaluated
 both at the community level and within species present on both paired
 aspects. The role of plasticity was examined in an experimental case
 study. We found that aspect is a strong determinant of phenology, with
 earlier flowering on warmer south-facing slopes. Aspect differences
 resulted in complementarity in timing of flowering resources across sites,
 as aspects that started flowering earlier also ended earlier.
 Complementarity between north and south aspects served to extend the
 flowering time of pollinator resources by an average of 4-8 days (8-15%),
 depending on the year. This extension can be attributed to both within
 species responses to aspect differences as well as species turnover.
 Flowering of L. gracilis genotypes was distinct across aspects,
 demonstrating that plasticity can drive the extension of flowering
 duration. Our findings indicate that heterogeneous topography can extend
 overall flowering time of pollinator resources, which may support
 pollinator biodiversity. Extension was most pronounced at the community
 level, which incorporates species turnover as well as plastic and
 genotypic differences within species.
description: Description of methods used for collection/generation of
 data: GrasslandPlotTemps2015_2018.csv
 Ibutton temperature data collected at Pepperwood Preserve in
 2015-2018. Paired grassland sites on North and South facing slopes.
 Temperatures were collected once per hour at 10cm below soil
 surface. Temperature and moisture were recorded at the
 plots to quantify microsite differences between aspects. Temperature was
 recorded at each aspect with an iButton (Thermochron, N=8) placed 10 cm
 below the soil surface and set to record every hour. Temperature
 measurements were taken from March 1<sup>st</sup> through June
 30<sup>th</sup> in all four years to compare aspect
 temperature differences during the flowering season. Additionally,
 temperature measurements were collected throughout the year in 2016 to
 capture full growing degree day accumulation curves.
 Phenology data collection methods: (for
 SpInfo.csv, PollResDates5.csv, SPECIESPollResDates.csv,
 LASPollResDates.csv) In each plot, we recorded
 flowering throughout the spring growing season (March-June) in 2015, 2016,
 2017 and 2018. Flowering phenology was observed for all pollinator
 resource species in the plots, including native and non-native species,
 annuals and perennials. Species status as a pollinator resource was
 identified by direct observations of animal visitation during the study,
 together with outside sources, including information provided by the
 Xerces society (Mader et al. 2011). Richness of pollinator resource
 species in each plot varied from 1 to 22 species over the entire season.
 Inflorescences in flower for each species in each plot were counted weekly
 to determine start, middle, and end flowering, as well as the length of
 the flowering season. For species with inflorescences that had more than
 one phenology stage present, an inflorescence was counted as flowering
 when at least 50% of it was in flower. Not all species were present in all
 sites, or on both slopes. Mader E, Shepherd M, Vaughan
 M, Hoffman Black S, LeBuhn. G (2011) The Xerces Society Guide to
 Attracting Native Pollinators: Protecting North America’s Bees and
 Butterflies. Storey Publishing, North Adams MA.
 Methods for data processing:
 PollResDates5.csv Raw flowering time data
 was collected as the number of inflorescences in flower for each species
 on each census date. Data was filtered to species that were determined to
 be pollinator resources. Phenology date columns
 calculation: The number of inflorescences on each census date in each plot
 was divided by the total number of inflorescences counted in the plot for
 the full season to determine proportional flowering on each date. This
 proportion was then summed to create a cumulative sum.
 Community flowering dates for pollinator resource species were
 calculated based on cumulative plot flowering over the season, as follows:
 start date as the date when 5% of the cumulative number of flowers in a
 plot (summed over the season) had been reached, mid-flowering date as the
 date when 50% flowering was reached, and end date as the date when 95% of
 flowering had been reached. Flowering duration was defined as the total
 number of days between start and end dates (when 5% and 95% flowering had
 been reached, respectively) for each plot.
 SPECIESPollResDates.csv Raw flowering time
 data was collected as the number of inflorescences in flower for each
 species on each census date. Data was filtered to species that were
 determined to be pollinator resources. Phenology date
 columns calculation: The number of inflorescences for each species on each
 census date in each plot was divided by the total number of inflorescences
 for that species counted in the plot for the full season to determine
 proportional flowering on each date. This proportion was then summed to
 create a cumulative sum for each species. Species
 flowering dates were defined as: start date as the date when 5% of the
 cumulative number of flowers in a plot (summed over the season) had been
 reached, mid-flowering date as the date when 50% flowering was reached,
 and end date as the date when 95% of flowering had been reached for each
 species. When flowering was only observed for a species on one survey
 date, the duration of flowering was calculated as 1 day (although it is
 likely that flowering occurred for two days or longer depending on the
 species). There were instances of gaps between flowering time on north and
 south aspects for individual species, and these were removed when
 calculating site flowering duration.
 LASPollResDates.csv Experimental plots of
 goldfields (<i>Lasthenia gracilis </i>(DC.) Greene) were set
 up just outside of main phenology plots in 2017 and 2018, with 3 subplots
 per aspect at each site (n=24). These 30x30 cm subplots were planted with
 30 seeds each, collected from two grassland locations on Pepperwood
 Preserve from 10 maternal lines (3 seeds per line per plot). Seeds were
 planted in the fall and marked with toothpicks to differentiate them from
 any other <i>Lasthenia</i> individuals occurring at the site.
 Flowering time was recorded for these subplots in the same way as the
 study phenology plots – with all open inflorescences from all individuals
 counted each week from March through June. Counts were conducted only on
 planted and marked <i>Lasthenia </i>individuals within each
 plot. Flowering dates were defined as: start date as
 the date when 5% of the cumulative number of flowers in a plot (summed
 over the season) had been reached, mid-flowering date as the date when 50%
 flowering was reached, and end date as the date when 95% of flowering had
 been reached for each experimental plot.
description: ReadMe file: PollinatorResources_readme.txt
 This PollinatorResources_readme.txt file was generated on
 2020-07-09 by Rachael L. Olliff-Yang GENERAL
 INFORMATION Data files and scripts related to the
 publication "Topographic heterogeneity lengthens the duration of
 pollinator resources" Author
 information: A. Principal Investigator Contact
 Information<br> Name: Rachael L. Olliff-Yang<br> Institution:
 University of California Berkeley<br> Address: 3040 Valley Life
 Sciences Building #3140<br> Email: rlolliff@berkeley.edu
 B. Co-investigator Contact Information<br> Name: David D.
 Ackerly<br> Institution: University of California Berkeley<br>
 Email: dackerly@berkeley.edu Years of data collection:
 2015-2018 Geographic location of data collection:
 Pepperwood Preserve (Sonoma County, California, USA 38.57˚ N, -122.68˚
 W) Funding sources that supported the collection of the
 data: Funding was provided by a Myrtle Wolf Scholarship
 (East Bay CNPS), a Natalie Hopkins Award (CNPS), a Mildred E. Mathias
 Research grant (UC NRS), and a UC Berkeley Integrative Biology graduate
 student research grant. Additional support was provided by the National
 Science Foundation Graduate Research Fellowship Grant (#1049702) (to
 R.L.O.-Y.). Code available at: https://github.com/rlolliff/Flowering-time-across-topography DATA & FILE OVERVIEW File List: BechtelMET_2014-2018.csv MoistureData.csv GrasslandPlotTemps2015_2018.csv AvMonthTemp_Full.csv PollResDates5.csv SPECIESPollResDates.csv LASPollResDates.csv SpInfo.csv Relationship between files: PollResDates5.csv, SPECIESPollResDates.csv, LASPollResDates.csv Were all calculated from SpInfo.csv raw phenology data. AvMonthTemp_Full.csv was calculated from GrasslandPlotTemps2015_2018.csv plot temperature data. Scripts available at https://github.com/rlolliff/Flowering-time-across-topography METHODOLOGICAL INFORMATION Description of methods used for collection/generation of data: GrasslandPlotTemps2015_2018.csv Ibutton temperature data collected at Pepperwood Preserve in 2015-2018. Paired grassland sites on North and South facing slopes. Temperatures were collected once per hour at 10cm below soil surface. Temperature and moisture were recorded at the plots to quantify microsite differences between aspects. Temperature was recorded at each aspect with an iButton (Thermochron, N=8) placed 10 cm below the soil surface and set to record every hour. Temperature measurements were taken from March 1<sup>st</sup> through June 30<sup>th</sup> in all four years to compare aspect temperature differences during the flowering season. Additionally, temperature measurements were collected throughout the year in 2016 to capture full growing degree day accumulation curves. Phenology data collection methods: (for SpInfo.csv, PollResDates5.csv, SPECIESPollResDates.csv, LASPollResDates.csv) In each plot, we recorded flowering throughout the spring growing season (March-June) in 2015, 2016, 2017 and 2018. Flowering phenology was observed for all pollinator resource species in the plots, including native and non-native species, annuals and perennials. Species status as a pollinator resource was identified by direct observations of animal visitation during the study, together with outside sources, including information provided by the Xerces society (Mader et al. 2011). Richness of pollinator resource species in each plot varied from 1 to 22 species over the entire season. Inflorescences in flower for each species in each plot were counted weekly to determine start, middle, and end flowering, as well as the length of the flowering season. For species with inflorescences that had more than one phenology stage present, an inflorescence was counted as flowering when at least 50% of it was in flower. Not all species were present in all sites, or on both slopes. Mader E, Shepherd M, Vaughan M, Hoffman Black S, LeBuhn. G (2011) The Xerces Society Guide to Attracting Native Pollinators: Protecting North America’s Bees and Butterflies. Storey Publishing, North Adams MA. Methods for data processing: PollResDates5.csv Raw flowering time data was collected as the number of inflorescences in flower for each species on each census date. Data was filtered to species that were determined to be pollinator resources. Phenology date columns calculation: The number of inflorescences on each census date in each plot was divided by the total number of inflorescences counted in the plot for the full season to determine proportional flowering on each date. This proportion was then summed to create a cumulative sum. Community flowering dates for pollinator resource species were calculated based on cumulative plot flowering over the season, as follows: start date as the date when 5% of the cumulative number of flowers in a plot (summed over the season) had been reached, mid-flowering date as the date when 50% flowering was reached, and end date as the date when 95% of flowering had been reached. Flowering duration was defined as the total number of days between start and end dates (when 5% and 95% flowering had been reached, respectively) for each plot. SPECIESPollResDates.csv Raw flowering time data was collected as the number of inflorescences in flower for each species on each census date. Data was filtered to species that were determined to be pollinator resources. Phenology date columns calculation: The number of inflorescences for each species on each census date in each plot was divided by the total number of inflorescences for that species counted in the plot for the full season to determine proportional flowering on each date. This proportion was then summed to create a cumulative sum for each species. Species flowering dates were defined as: start date as the date when 5% of the cumulative number of flowers in a plot (summed over the season) had been reached, mid-flowering date as the date when 50% flowering was reached, and end date as the date when 95% of flowering had been reached for each species. When flowering was only observed for a species on one survey date, the duration of flowering was calculated as 1 day (although it is likely that flowering occurred for two days or longer depending on the species). There were instances of gaps between flowering time on north and south aspects for individual species, and these were removed when calculating site flowering duration. LASPollResDates.csv Experimental plots of goldfields (<i>Lasthenia gracilis </i>(DC.) Greene) were set up just outside of main phenology plots in 2017 and 2018, with 3 subplots per aspect at each site (n=24). These 30x30 cm subplots were planted with 30 seeds each, collected from two grassland locations on Pepperwood Preserve from 10 maternal lines (3 seeds per line per plot). Seeds were planted in the fall and marked with toothpicks to differentiate them from any other <i>Lasthenia</i> individuals occurring at the site. Flowering time was recorded for these subplots in the same way as the study phenology plots – with all open inflorescences from all individuals counted each week from March through June. Counts were conducted only on planted and marked <i>Lasthenia </i>individuals within each plot. Flowering dates were defined as: start date as the date when 5% of the cumulative number of flowers in a plot (summed over the season) had been reached, mid-flowering date as the date when 50% flowering was reached, and end date as the date when 95% of flowering had been reached for each experimental plot. DATA-SPECIFIC INFORMATION FOR: BechtelMET_2014-2018.csv Number of variables: 10 Number of cases/rows: 175053 Variable List: “timestamp” - mm/dd/yyyy hh:mm “DOY” = day of year (# of days since Jan1st) “WDOY” – day of water year (#days since October 1<sup>st</sup>) “YEAR” – year of collection “the records” = record # (for reference) “airTemp” = degrees C “rainHun15min” = Number of tips 1tip =0.01" of precip “rainInchYear” = Cummulative sum of precip for water year beginning OCT 1 (inches) “rainInch15min” = Number of tips / 100 (inches) Bechtel weather station and rain gauge data collected every 15 minutes.   LIST OF OMMITTED DATA TIMESTAMP START TIMESTAMP END FIELD REASON 12/9/15 16:15 2/2/16 17:15 AirTC_Avg Sensor malfunction and removal for calibration 12/9/15 16:15 2/2/16 17:15 RH_Avg Sensor malfunction and removal for calibration. Bad values (over 100 preced this period, RH was out of calibration) 10/9/17 4:45 1/5/18 10:30 TE 525 15min Rain, ("rainHun15min", "raininchYear", "rainInch15min", "rainInch15minForm") Tubbs Fire knocked over rain gauge, not reinstalled until after recalibration. Nearby gauge recorded 8.6 inches during this time 12/5/17 7:15 1/5/18 10:30 All Sensors sent in for recalibrations. 1/8/18 11:00 1/8/18 11:00 barometricPressure dropped out unknown but fell from ~750mmHg to 20-50mmHg for one 15minute average 4/13/18 11:00 4/13/18 11:00 barometricPressure dropped out unknown but fell from ~750mmHg to 20-50mmHg for one 15minute average 6/11/18 8:00 6/11/18 8:00 barometricPressure dropped out unknown but fell from ~750mmHg to 20-50mmHg for one 15minute average 8/14/18 9:30 8/14/18 9:30 barometricPressure dropped out Logger powered off as hard reboot to reestablish internet connection to DC. -RF   DATA-SPECIFIC INFORMATION FOR: MoistureData.csv Number of variables: 8 Number of cases/rows: 163 Variable List: “DATE” – Date of soil moisture measurement; mm/dd/yy “DOY” – day of year (# of days since Jan1st) “YEAR” – year of collection “PLOT”- indicates site (1<sup>st</sup> 2 letters), aspect (3<sup>rd</sup> letter), and replicate (1, 2, or 3). “SITE” – Location of hill with paired aspects (BH = Bechtel House, DP= Double ponds, FH=Hill 1521, TP = Turtle ponds, TH = Three Tree Hill. “ASPECT” -aspect of plot (N = North, S= South) “PERCENTVWC” –Volumetric Water Content (%) “Microsiemens” – Electrical conductivity (µS) DATA-SPECIFIC INFORMATION FOR: GrasslandPlotTemps2015_2018.csv Number of variables: 9 Number of cases/rows: 155780 Variable List: “PLOT”- indicates site (1<sup>st</sup> 2 letters) and aspect (3<sup>rd</sup> letter) where temperature data was collected. This does not correspond to individual replicate “plots” from related data in this repository, only to the site and aspect (since temperature data was taken from a central point between the three plots.) “DateTime” – date and time of temperature measurement;mm/dd/yyyy hh:mm “DOY” – day of year (# of days since Jan1st) “WDOY” – day of water year (#days since October 1<sup>st</sup>) “YEAR” – year of collection “WYEAR” – water year of collection (October-September) “TEMP” – temperature measurement (˚C) “SerialNo” – Serial number of ibutton for testing accuracy and comparison “DataType” – Plot (temperature taken at plot), Test (temperatures taken before/after placement in plot, in same location as other ibuttons for testing serial number differences in temperature readings), PlotBAD (ibutton data taken at plot but disturbed/displaced/unearthed by animal activity) Ibutton temperature data collected at Pepperwood Preserve in 2015-2018. Paired grassland sites on North and South facing slopes. Temperatures were collected once per hour at 10cm below soil surface. DATA-SPECIFIC INFORMATION FOR: AvMonthTemp_Full Number of variables: 17 Number of cases/rows: Variable List: “YEAR” – year of data collection (2015, 2016, 2017, 2018) “PLOT”- indicates site (1<sup>st</sup> 2 letters) and aspect (3<sup>rd</sup> letter) where temperature data was collected. This does not correspond to individual replicate “plots” from related data in this repository, only to the site and aspect (since temperature data was taken from a central point between the three plots.) “TempJan” – average temperature (˚C) 10cm below soil surface in January “TempFeb”– average temperature (˚C) 10cm below soil surface in February “TempMar”– average temperature (˚C) 10cm below soil surface in March “ExtrMarch” – extrapolated March temperature data (˚C)for plots with missing data. Calculated as the average march temperature at that plot in other years and adjusted by the average difference in march air temperature data. “TempApril”– average temperature (˚C) 10cm below soil surface in April “ExtrApril “– extrapolated April temperature data (˚C)for plots with missing data. Calculated as the average April temperature at that plot in other years and adjusted by the average difference in April air temperature data. “TempMay”– average temperature (˚C) 10cm below soil surface in May “ExtrMay” – extrapolated May temperature data (˚C)for plots with missing data. Calculated as the average May temperature at that plot in other years and adjusted by the average difference in May air temperature data. “TempJun”– average temperature (˚C) 10cm below soil surface in June “TempJul”– average temperature (˚C) 10cm below soil surface in July “TempAug”– average temperature (˚C) 10cm below soil surface in August “TempSep”– average temperature (˚C) 10cm below soil surface in Septemeber “TempOct”– average temperature (˚C) 10cm below soil surface in October “TempNov”– average temperature (˚C) 10cm below soil surface in November “TempDec”– average temperature (˚C) 10cm below soil surface in December Data was averaged from hourly measurements, catalogued in GrasslandPlotTemps2015_2018.csv DATA-SPECIFIC INFORMATION FOR: PollResDates5.csv Number of variables: 8 Number of cases/rows: 97 Variable List: “YEAR” – year of data collection (2015, 2016, 2017, 2018) “ASPECT” -aspect of plot (N = North, S= South) “SITE” – Location of hill with paired aspects (BH = Bechtel House, DP= Double ponds, FH=Hill 1521, TP = Turtle ponds, TH = Three Tree Hill). TH was only included in 2016 & 2017 and was not included in ANOVA analyses. “PLOT” – Six per site, Three plots on each aspect. 1<sup>st</sup> two letters (e.g. BH) indicate site, 3<sup>rd</sup> (e.g. N) indicates aspect, Number at end (1, 2, or 3) indicates replicate. “Start” day of year of 5% flowering. “Mid” or “Peak” day of year of 50% flowering, “End” the day of year of 95% flowering. “length” = number of days from “start” (5% flowering) to “end” (95% flowering) dates Raw flowering time data was collected as the number of inflorescences in flower for each species on each census date. Data for this paper was filtered to species that were determined to be pollinator resources. Phenology date columns calculation: The number of inflorescences on each census date in each plot was divided by the total number of inflorescences counted in the plot for the full season to determine proportional flowering on each date. This proportion was then summed to create a cumulative sum. DATA-SPECIFIC INFORMATION FOR: SPECIESPollResDates.csv Number of variables: 9 Number of cases/rows: 995 Variable List: “YEAR” – year of data collection (2015, 2016, 2017, 2018) “ASPECT” -aspect of plot (N = North, S= South) “SITE” – Location of hill with paired aspects (BH = Bechtel House, DP= Double ponds, FH=Hill 1521, TP = Turtle ponds, TH = Three Tree Hill). TH was only included in 2016 & 2017 and was not included in ANOVA analyses. “PLOT” – Six per site, Three plots on each aspect. 1<sup>st</sup> two letters (e.g. BH) indicate site, 3<sup>rd</sup> (e.g. N) indicates aspect, Number at end (1, 2, or 3) indicates replicate. “NAME” –Species binomial ““Start” day of year of 5% flowering. “Mid” or “Peak” day of year of 50% flowering, “End” the day of year of 95% flowering. “length” = number of days from “start” (5% flowering) to “end” (95% flowering) dates Phenology date columns calculation: The number of inflorescences for each species on each census date in each plot was divided by the total number of inflorescences for that species counted in the plot for the full season to determine proportional flowering on each date. This proportion was then summed to create a cumulative sum for each species. Raw flowering time data was collected as the number of inflorescences in flower for each species on each census date. Data for this paper was filtered to species that were determined to be pollinator resources. DATA-SPECIFIC INFORMATION FOR: LASPollResDates.csv Number of variables: 8 Number of cases/rows: 38 Variable List: “YEAR” – year of experiment data collection (2017, 2018) “ASPECT” - aspect of plot (N = North, S= South) “SITE” – Location of hill with paired aspects (BH = Bechtel House, DP= Double ponds, FH=Hill 1521, TP = Turtle ponds, TH = Three Tree Hill). TH was only included in 2016 & 2017 and was not included in ANOVA analyses. “PLOT” – Six per site, Three plots on each aspect. 1<sup>st</sup> two letters (e.g. BH) indicate site, 3<sup>rd</sup> (e.g. N) indicates aspect, Number at end (1, 2, or 3) indicates replicate. “Start” day of year of 5% flowering. “Mid” or “Peak” day of year of 50% flowering, “End” the day of year of 95% flowering. “length” = number of days from “start” (5% flowering) to “end” (95% flowering) day of year Raw flowering time data was collected as the number of inflorescences in flower for <i>Lasthenia gracilis</i> in experimental plots on each census date. Phenology date columns calculation: The number of <i>Lasthenia gracilis</i> inflorescences in experimental plots on each census date in each plot was divided by the total number of inflorescences counted in the plot for the full season to determine proportional flowering on each date. This proportion was then summed to create a cumulative sum. DATA-SPECIFIC INFORMATION FOR: SpInfo.csv Number of variables: 22 Number of cases/rows: 17022 Variable List: "SP_CODE" – six (or nine) letter unique species code (1<sup>st</sup> 3 letters of genus and 1<sup>st</sup> 3 letters of specific epithet). Nine letter codes are used in cases where a six letter code would be identical to another species already in the dataset. "DATE" - mm/dd/yy “YEAR” – year of experiment data collection (2017, 2018) “DOY” – day of year (# of days since Jan1st) “PLOT” – Six per site, Three plots on each aspect. 1<sup>st</sup> two letters (e.g. BH) indicate site, 3<sup>rd</sup> (e.g. N) indicates aspect, Number at end (1, 2, or 3) indicates replicate. “SITE” – Location of hill with paired aspects (BH = Bechtel House, DP= Double ponds, FH=Hill 1521, TP = Turtle ponds, TH = Three Tree Hill). TH was only included in 2016 & 2017 and was not included in ANOVA analyses. “ASPECT” - aspect of plot (N = North, S= South) "PHENO"- all reproductive phases observed "MAIN_PHENO"- main reproductive phase "INFL" - number of total inflorescences in bud, flower or fruit "FLS" - number of inflorescences in flower "FRT" - number of inflorescences in fruit “NAME” – Species binomial "LIFESPAN"- A = annual, P= perennial "TYPE" – F=Forb, G=Grass "NATIVE"- Y = yes, N= No "PollinatorResource"- pollinator resource Y = yes, N= No "BEE"- Bee resource Y = yes, N= No "BUTTERFLY"- butterfly resource Y = yes, N= No "FLY"= fly resource Y = yes, N= No "MOTHS" – moth resource Y = yes, N= No "BEETLES"- beetle resource Y = yes, N= No Raw phenology data for all species in plots, collected as the number of inflorescences in flower and fruit. Species status as a pollinator resource was identified by direct observations of animal visitation during the study, together with outside sources, including information provided by the Xerces society (Mader et al. 2011). Mader E, Shepherd M, Vaughan M, Hoffman Black S, LeBuhn. G (2011) The Xerces Society Guide to Attracting Native Pollinators: Protecting North America’s Bees and Butterflies. Storey Publishing, North Adams MA.