DOI: 10.5061/dryad.0k6djhbb8
title: Selection maintains floral color polymorphism in the scarlet paintbrush, <em>Castilleja coccinea</em>, reflecting combined ecological factors

description: Premise: Evolutionary theory predicts polymorphism should be rare;
 however, variation in floral color is common, often attributed to drift,
 plasticity, or variable selection. Examining floral color polymorphism
 both within contact zones and across a species’ range can reveal
 mechanisms maintaining this variation. Here, we used a multistep approach
 to investigate spatially heterogeneous variation in floral bract color in
 Castilleja coccinea. Methods: We compared frequencies of color morphs,
 floral morphology, fitness, and genetic structure in regional populations
 and a common garden. Next, we examined habitat differences, including
 plant communities and edaphic factors, as potential drivers of variation.
 Lastly, we leveraged herbarium and iNaturalist occurrence data to
 investigate whether patterns were consistent at the landscape scale. Key
 results: Bract color in C. coccinea is genetically heritable, with yellow
 dominant over red, and is under selection. Populations are predominantly
 monomorphic, with color distance showing no correlation to genetic or
 geographic distance, despite significant genetic isolation-by-distance.
 Yellow morphs are associated with open wetlands, while red morphs occur at
 drier sites associated with nearby tree cover. Red morphs demonstrated
 lower fitness in a common garden, suggesting tradeoffs associated with
 pleiotropic effects of adaptation to drier soil conditions. Conclusions:
 Differences in floral color between morphs are consistent with
 diversification associated with a shift in ecological niche. We identified
 variation in edaphic and habitat conditions as probable drivers of
 divergence in floral color. Additionally, variation in other floral traits
 suggests a combined role of pollinators and habitat differences acting in
 concert to maintain distinct floral color morphs.
description: # Selection maintains floral color polymorphism in the scarlet paintbrush
 *Castilleja coccinea*, reflecting combined ecological factors Dataset DOI:
 [10.5061/dryad.0k6djhbb8](10.5061/dryad.0k6djhbb8) ## Description of the
 data and file structure In this study, we sought to characterize and
 determine the drivers of floral bract color polymorphism in *Castilleja
 coccinea*, both in a contact zone of morphotypes and across the range of
 the species. We focus on four primary objectives: **1)** establishing that
 this variation is the result of selection; **2)** determining if the trait
 is genetically inherited and associated with fitness differences; **3)**
 identifying potential abiotic or biotic agents of selection that may be
 driving these patterns; and **4)** determining if these patterns are
 generalizable across the range of the species. To accomplish this, we have
 completed field studies of floral traits and habitat composition, a common
 garden experiment, an analysis of genetic structure, and a landscape-scale
 analysis of herbarium and community science data.  The data files are
 organized into four directories: 1) Landscape scale data
 (Landscape_data.zip), 2) Morphology (Morphology_data.zip), 3) Habitat
 (Habitat_data.zip), and 4) Population genomics data
 (Genetics_denovo_data.zip).  Note: NA in this dataset indicates data that
 is not available. ### Files and variables ### Landscape scale data *All
 contained in the Landscape_data.zip directory* Note: GPS coordinates
 (latitude and longitude) of GBIF observations across the species’ range in
 this directory have been rounded to the nearest 0.1° to protect
 potentailly sensitive populations. For access to precise location data,
 please contact the corresponding author. The only exceptions are
 populations in Illinois, where *Castilleja coccinea* is considered secure
 or has no status rank. #### File: caco_all_map_rounded.csv
 **Description:** Derived datset of herbarium and iNaturalist observations
 used to create the rangewide map figure (Figure 1A). This file contains 6
 columns and 3701 rows (obs) of data. ##### Variables * ID: unique
 identifier for each observation, either the iNaturalist ID number, or the
 herbarium ID * latitude: north–south position of a point * longitude:
 east–west position of a point * color: describes the bract color of the
 observation as Red or Yellow * Record_type: describes the origin of data
 as either from iNaturalist, herbarium or field collected data * name:
 scientific name of species (Castilleja coccinea) #### File:
 Anna_samples_gps.csv **Description:** Field sites used to create the inset
 map figure (Figure 1B). This file contains 7 columns and 11 rows (obs) of
 data. ##### Variables * Site: abbreviated site name relating to geographic
 location of observation IB2 = Illinois Beach 2, MW = Newark Road, = Pine
 Station, SP = Shaw Prairie, IB1 = Illinois Beach 1, DP = Dropseed Prairie,
 HP = Hoosier Prairie, MC = Meissner-Corron, NR = Newark Road, SR = Sand
 Ridge, GM = Gensburg-Markham * lat: north–south position of a point *
 long: east–west position of a point * color: describes the bract color of
 the observation as Red or Yellow * site_code: unique alpha numerical
 description of site, R = red site, Y = yellow site, P = polymorphic site
 #### File: color_complete_rounded.csv **Description:** Cleaned and
 filtered output of the ColorSelector pipeline (Luong et al. 2023). This
 file contains 35 columns and 2447 rows (obs) of data. ##### Variables *
 iNatID: unique identifier of observation derived from
 ihttps://[www.inaturalist.org/](http://www.inaturalist.org/) * species:
 species name * long: east–west position of a point * lat: north–south
 position of a point * year: year of observation * month: month of
 observation * day: month of observation * Nsamples: number of pixels
 sampled for one observation using the ColorSelector R shiny applicaiton *
 Rmean: mean red value of sampled pixels in obeservation * Rmedian: median
 of red value of sampled pixels in obeservation * Rmax: maximum of red
 value for sampled pixels * Rmin: minimum of red value for sampled pixels *
 Rvariance: variance of red values for samples pixels * Rstdev: standard
 deviation of red values for sampled pixels * Gmean: mean green value of
 sampled pixels in obeservation * Gmedian: median of green value of sampled
 pixels in observation * Gmax: maximum of green value for sampled pixels *
 Gmin: minimum of green value for sampled pixels * Gvariance: variance of
 green values for samples pixels * Gstdev: standard deviation of green
 values for sampled pixels * Bmean: mean blue value of sampled pixels in
 observation * Bmedian: median of blue value of sampled pixels in
 observation * Bmax: maximum of blue value for sampled pixels * Bmin:
 minimum of blue value for sampled pixels * Bvariance: variance of blue
 values for sample pixels * Bstdev: standard blue of green values for
 sampled pixels * poly: if the observation represents a polymorphic
 population. values "Y" = yes, "N" = No * RGB_vector:
 Bract color RGB values were transformed into a single vector value by
 treating each color as a point in n-dimensional space and determining the
 Euclidean distance of each point from the origin. * Hue: hue value of
 observation (from HSV spectrum) * Saturation: saturation value of
 observation (from HSV spectrum) * Value: value of observation (from HSV
 spectrum) * Category: bract color category. values "Red" or
 "Yellow" or "Orange" * Category_binary: binary bract
 color category "Red" or "Yellow" * Fill: hex code of
 hue value * Adjusted_hue: hue value adjusted from a #### File:
 anna_confirmed_final_rounded.csv **Description:** Confirmed and filtered
 herbarium specimen information. This file contains 23 columns and 1322
 rows (obs) of data. ##### Variables * ID: Unique identifier for each
 herbarium record * Accession.ID: Accession or catalog identifier for the
 sample * Anna.Data: Reference to data associated with Anna * Duplicate:
 Indicates duplicate samples * Herbarium.ID: Herbarium specieman * lat: GPS
 coordinates (latitude) * long: GPS coordinates (longitude) * Accuracy:
 Accuracy of the GPS coordinates (character) * County: County where the
 sample is located * State: State where the sample is located *
 Collection.Month: Month of sample collection * Collection.Day: Day of the
 month for sample collection * Collection.Yr: Year of sample collection *
 Day.Month: Date in day/month format * Julian.Days: Day of the year of
 collection (Julian date) * Accuracy.of.date: Precision of collection date
 (character) * Yellow: Indicates presence of yellow color or label
 description (TRUE or FALSE) * Confirmed: Indicates confirmation status of
 bract color (TRUE or FALSE) * Color.Notes: Additional notes on color from
 herbarium label * Location: Location information for the sample * Habitat:
 Habitat type where the sample was collected * Incomplete: Indicates
 incomplete data on label * color: Color classification of the sample
 (factor; yellow or red) #### File: COMPLETE_Herbarium_rounded.csv
 **Description:** Herbarium specimen data file - unfiltered. This file
 contains 31 columns and 2883 rows (obs) of data. ##### Variables * ID:
 Unique identifier for each herbarium record * Accession.ID: Accession or
 catalog identifier for the sample * Anna.Data: Reference to data
 associated with Anna (TRUE or FALSE) * Duplicate: Indicates duplicate
 herbarium collections * Herbarium.ID: Herbarium name * GPS.North: GPS
 coordinates (latitude) * GPS.West: GPS coordinates (longitude) * Accuracy:
 Accuracy of the GPS coordinates (character) * County: County where the
 sample is located * State: State where the sample is located *
 Collection.Month: Month of sample collection * Collection.Day: Day of the
 month for sample collection * Collection.Yr: Year of sample collection *
 Day.Month: Date in day/month format * Julian.Days: Day of the year (Julian
 date) * Accuracy.of.date: Precision of collection date * Yellow: Indicates
 presence of yellow color or label description (TRUE or FALSE) * Confirmed:
 Indicates confirmation status of bract color (TRUE or FALSE) *
 Color.Notes: Additional notes on color from herbarium label * Location:
 Location information for the sample * Habitat: Habitat type where the
 sample was collected * PLSS.data: Public Land Survey System (PLSS) data *
 TSR: Township, section, and range (PLSS data) * Incomplete: Indicates
 incomplete data on label * PLSS.notes: Additional notes related to PLSS
 data * Notes: General notes about the sample * GPS.North.original:
 Original GPS latitude before adjustments * GPS.West.original: Original GPS
 longitude before adjustments * GPS.PLSS: GPS coordinates converted to PLSS
 format * GPS.Google.Earth: GPS coordinates from Google Earth *
 Acc.to.Sect: Accession to section reference #### File:
 CACO_colors_rounded.csv **Description:** Output of the ColorSelector
 pipeline averaged by observation - unfiltered. This file contains 19
 columns and 2525 rows (obs) of data. ##### Variables * iNatID: Unique
 identifier of observation derived from
 ihttps://[www.inaturalist.org/](http://www.inaturalist.org/) * species:
 species name * long: east–west position of a point * lat: north–south
 position of a point * year: year of observation * month: month of
 observation * day: month of observation * Nsamples: number of pixels
 sampled for one observation using the ColorSelector R Shiny application *
 R.mean: Mean red intensity value in the RGB color model * R.median: Median
 red intensity value in the RGB color model * R.max: Maximum red intensity
 value in the RGB color model * R.min: Minimum red intensity value in the
 RGB color model * R.variance: Variance of red intensity values in the RGB
 color model * R.stdev: Standard deviation of red intensity values in the
 RGB model * G.mean: Mean green intensity value in the RGB color model *
 G.median: Median green intensity value in the RGB color model * G.max:
 Maximum green intensity value in the RGB color model * G.min: Minimum
 green intensity value in the RGB color model * G.variance: Variance of
 green intensity values in the RGB color model * G.stdev: Standard
 deviation of green intensity values in the RGB model * B.mean: Mean blue
 intensity value in the RGB color model * B.median: Median blue intensity
 value in the RGB color model * B.max: Maximum blue intensity value in the
 RGB color model * B.min: Minimum blue intensity value in the RGB color
 model * B.variance: Variance of blue intensity values in the RGB color
 model * B.stdev: Standard deviation of blue intensity values in the RGB
 model * poly: Polymorphic site, if site had more than one floral morph
 visible in iNat image ### Morphology data *All files contained in
 the Morphology_data.zip directory.* #### File:
 CACO_ccHSV_field_20250506.csv **Description:** Bract color, bract
 measurements, floral measurements, RGB, HSV values for individual plants,
 and proportion of color by site This file contains 36 columns and 448 rows
 (obs) of data. ##### Variables * Site Name: site name; geographic location
 of observation (Text) Illinois Beach 2, Newark Road, Pine Station, Shaw
 Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * Plan ID:
 unique numerical ID for each individual plant with decimal indicating the
 flower measured on each plant * Bract.Color: alpha numeric Royal
 Horticulture Society code for color * ID: unique numerical ID for each
 flower * Plant ID: unique numerical ID for each individual plant * Tag No:
 unique numerical ID for each individual plant * Bract.Length: length of
 bract from bact base to lobe tips (mm) * Bract.Base: length of bract base
 to lobe base (mm) * Bract.Width: width of bract at widest point (mm) *
 Bract.Lobe: count of number of bract lobes * Calyx.Outer: length of outer
 calyx segment (mm) * Calyx.Inner: length of inner calyx segment (mm) *
 Corolla.Tube.Length: length of corrolla tube (mm) * Corolla.Galea.Length:
 length of corrolla galea (mm) * Corolla.Tube.Width: width of corrolla tube
 (mm) * Stamen.Length: stamen length (mm) * Style.Length: style length (mm)
 * Herkogamy: separation of anthers and stigma (stamen length - stigma
 length) * R: red value in RGB color space * G: green value in RGB color
 space * B: blue value in RGB color space * hex: hexcode from RGB color
 vector * Hue: hue value in HSV colorspace * Saturation: saturation value
 in HSV colorspace * Value: value in HSV color space * Category: bract
 color category. values "Red" or "Yellow" or
 "Orange" * Fill: hexcode from hue value * Adjusted_Hue: hue
 value corrected for circularity * observation: value of 1 * Category2:
 bract color category values "Red" or "Yellow" *
 SitePropRed: proportion of plants that were red * SitePropYellow:
 proportion of plants that were yellow * Site.Color: dominant site color as
 "Red" or "Yellow" * Site.ColorFine: dominant site
 color as "Red" or "Yellow" or "Yellow_mixed"
 * morphSite: individual plant color with sites color (eg. Red - Yellow) *
 pch_ind: shape values as 21 (circle) or 22 (square) #### File:
 CACO_cgHSV_cg_20250506.csv **Description:** Bract color, bract
 measurements, floral measurements, RGB, HSV values for individual plants,
 and proportion of color by (maternal) site This file contains 37 columns
 and 841 rows (obs) of data. ##### Variables * Site Name: site name;
 geographic location of observation (Text) Illinois Beach 2, Newark Road,
 Pine Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie, Hoosier
 Prairie, Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham *
 Plant ID: unique numerical ID for each individual plant with decimal
 indicating the flower measured on each plant * Maternal Bract Color: alpha
 numeric Royal Horticulture Society code for maternal bract color * Common
 garden ID: umerical ID for common garden * RHS.Code: alpha numeric Royal
 Horticulture Society code for plant color * ID1: unique numerical ID for
 each individual plant * CGID: umerical ID for common garden * BractLength:
 length of bract from bact base to lobe tips * BractBase: length of bract
 base to lobe base * BractWidth: width of bract at widest pointt *
 Bract.Lobe: count of number of bract lobes * Calyx.Outer: length of outer
 calyx segment (mm) * Calyx.Inner: length of inner calyx segment (mm) *
 Corolla.Tube.Length: length of corrolla tube (mm) * Corolla.Galea.Length:
 length of corrolla galea (mm) * Corolla.Tube.Width: width of corrolla tube
 (mm) * Stamen.Length: stamen length (mm) * Style.Length: style length (mm)
 * Herkogamy: separation of anthers and stigma (stamen length - stigma
 length)) * R: red value in RGB color space * G: green value in RGB color
 space * B: blue value in RGB color space * hex: hexcode from RGB color
 vector * Hue: hue value in HSV colorspace * Saturation: saturation value
 in HSV colorspace * Value: value in HSV color space * Adjusted_Hue: hue
 value corrected for circularity * Category: bract color category. values
 "Red" or "Yellow" or "Orange" * Fill:
 hexcode from hue value * Category2: bract color category values
 "Red" or "Yellow" * observation: observation *
 SitePropYellow: proportion of plants that were yellow * Site.Color:
 dominant site color as "Red" or "Yellow" *
 Site.ColorFine: dominant site color as "Red" or
 "Yellow" or "Yellow_mixed" * pch_ind: shape values as
 21 (circle) or 22 (square) * nmds1m: nmds axis 1 position * nmds2m: nmds
 axis 2 position #### File: CACO_CGcolorOffspring_collapsed_20241115.csv
 **Description:** Offspring of controlled cross experiment. This file
 contains 48 columns and 24 rows (obs) of data. ##### Variables *
 Site.Name: site name; geographic location of observation (Text) Illinois
 Beach 2, Newark Road, Pine Station, Shaw Prairie, llinois Beach 1,
 Dropseed Prairie, Hoosier Prairie, Meissner-Corron, Newark Road, Sand
 Ridge, Gensburg-Markham * MaternalBractColor: alpha numeric Royal
 Horticulture Society code for maternal bract color * MaternalColCatBroad:
 bract color category values "Red" or "Yellow" * count:
 count of offsping * os_2D: Offspring RHS Color: 2D * os_3A: Offspring RHS
 Color: 3A * os_3B: Offspring RHS Color: 3B * os_3C: Offspring RHS Color:
 3C * os_4A: Offspring RHS Color: 4A * os_4B: Offspring RHS Color: 4B *
 os_4C: Offspring RHS Color: 4C * os_5A: Offspring RHS Color: 5A * os_5B:
 Offspring RHS Color: 5B * os_6A: Offspring RHS Color: 6A * os_7A:
 Offspring RHS Color: 7A * os_7B: Offspring RHS Color: 7B * os_8A:
 Offspring RHS Color: 8A * os_9A: Offspring RHS Color: 9A * os_9B:
 Offspring RHS Color: 9B * os_10A: Offspring RHS Color: 10A * os_12A:
 Offspring RHS Color: 12A * os_12B: Offspring RHS Color: 12B * os_13A:
 Offspring RHS Color: 13A * os_13B: Offspring RHS Color: 13B * os_21A:
 Offspring RHS Color: 21A * os_22A: Offspring RHS Color: 22A * os_22B:
 Offspring RHS Color: 22B * os_23A: Offspring RHS Color: 23A * os_25A:
 Offspring RHS Color: 25A * os_25B: Offspring RHS Color: 25B * os_N25A:
 Offspring RHS Color: N25A * os_N25B: Offspring RHS Color: N25B * os_26A:
 Offspring RHS Color: 26A * os_28A: Offspring RHS Color: 28A * os_30A:
 Offspring RHS Color: 30A * os_30B: Offspring RHS Color: 30B * os_N30A:
 Offspring RHS Color: N30A * os_N30B: Offspring RHS Color: N30B * os_N30C:
 Offspring RHS Color: N30C * os_31A: Offspring RHS Color: 31A * os_32A:
 Offspring RHS Color: 32A * os_33A: Offspring RHS Color: 33A * os_33B:
 Offspring RHS Color: 33B * os_35A: Offspring RHS Color: 35A * os_39B:
 Offspring RHS Color: 39B * os_N39B: Offspring RHS Color: N39B * os_yellow:
 offspring yellow * os_red: offsping red #### File: CG_bract_only.csv
 **Description:** Bract measurements (length, width, number etc) of
 individual plants in the common garden. This file contains 13 columns and
 442 rows (obs) of data. ##### Variables * Site.Name: site name; geographic
 location of observation (Text) Illinois Beach 2, Newark Road, Pine
 Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * ColorS:
 dominant site color as "Yellow" or "Orange" *
 MplantID: maternal plant unique numerical ID * ColorM.RHS.: alpha numeric
 Royal Horticulture Society code for maternal bract color * ColorM:
 maternal plant color as "Yellow" or "Orange" * CGID:
 unique plant ID number in common garden * ColorP: color of the individual
 plant as "Yellow" or "Orange" * ColorP..RHS.: alpha
 numeric Royal Horticulture Society code for individual bract color *
 Bract.Length: length of bract from bact base to lobe tips * Bract.Base:
 length of bract base to lobe base * Bract.Width: width of bract at widest
 pointt * BractNo: count of number of bract lobes * ColorSP: color of the
 site (S) with color of the individual plant (P) #### File: CG_floral.csv
 **Description:** Floral measurements (Calyx, corolla, stamen, style etc)
 of individual plants in the common garden. This file contains columns 17
 columns and 431 rows (obs) of data. ##### Variables * Site.Name: site
 name; geographic location of observation (Text) Illinois Beach 2, Newark
 Road, Pine Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie,
 Hoosier Prairie, Meissner-Corron, Newark Road, Sand Ridge,
 Gensburg-Markham * ColorS: dominant site color as "Yellow" or
 "Orange" * MPlant.ID: maternal plant unique numerical ID *
 ColorM..RHS.: alpha numeric Royal Horticulture Society code for maternal
 bract color * ColorM: maternal plant color as "Yellow" or
 "Orange" * CGID: unique plant ID number in common garden *
 ColorP: color of the individual plant as "Yellow" or
 "Orange" * ColorP..RHS.: alpha numeric Royal Horticulture
 Society code for individual bract color * Calyx.Outer: length of outer
 calyx segment (mm) * Calyx.Inner: length of inner calyx segment (mm) *
 Corolla.Tube.Length: length of corrolla tube (mm) * Corolla.Galea.Length:
 length of corrolla galea (mm) * Corolla.Tube.Width: width of corrolla tube
 (mm) * Stamen.Length: stamen length (mm) * Style.Length: style length (mm)
 * Herkogamy: separation of anthers and stigma (stamen length - stigma
 length)) * ColorSP: color of the site (S) with color of the individual
 plant (P) #### File: Field_bract_only.csv **Description:** Bract
 measurements (length, width, number etc) of individual plants in the
 field. This file contains 10 columns and 336 rows (obs) of data. #####
 Variables * site name: site name (Text); geographic location of
 observation Illinois Beach 2, Newark Road, Pine Station, Shaw Prairie,
 llinois Beach 1, Dropseed Prairie, Hoosier Prairie, Meissner-Corron,
 Newark Road, Sand Ridge, Gensburg-Markham * ColorS: dominant bract color
 of site grouped into two categories (Yellow or Orange) * PlantID: unique
 numerical ID for each individual plant * ColorID: alpha numeric Royal
 Horticulture Society code for color * ColorP: plant bract color grouped
 into two categories (Yellow or Orange) * Bract length : length of floral
 bract measured from base to tip (mm) * Bract Base: width of floral bract
 base (mm) * Bract Width: width of floral bract at its widest point (mm) *
 BractNo: number of bracts (integer) * ColorSP: combination of site color
 (ColorS = dominant bract color of site grouped into two categories (Yellow
 or Orange)) and plant color (ColorP = individual plant bract color grouped
 into two categories (Yellow or Orange)) #### File: Field_floral.csv
 **Description:** Floral measurements (calyx, corolla, stamen, style etc)
 of individual plants in the field. This file contains 14 columns and 478
 rows (obs) of data. ##### Variables * site name: site name; geographic
 location of observation (Text) Illinois Beach 2, Newark Road, Pine
 Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * ColorS:
 dominant bract color of site grouped into two categories (Yellow or
 Orange) * Plant ID: unique numerical ID for each individual plant * Color
 (RHS): alpha numeric Royal Horticulture Society code for color * ColorP:
 plant bract color grouped into two categories (Yellow or Orange) * Calyx
 Outer: length of outer calyx segment (mm) * Calyx Inner: length of inner
 calyx segment (mm) * Corrolla Tube: length of corrolla tube (mm) *
 BractNo: number of bracts (integer) * Corolla Galea Length: length of
 corrolla galea (mm) * Corolla Tube Width: width of corrolla tube (mm) *
 Stamen Length: stamen length (mm) * Style Length: style length (mm) *
 Herkogamy: separation of anthers and stigma (stamen length - stigma
 length)) * ColorSP: combination of site color (ColorS = dominant bract
 color of site grouped into two categories (Yellow or Orange)) and plant
 color (ColorPwith individual plant bract color grouped into two categories
 (Yellow or Orange)) #### File: Fruit_flower_CG.csv **Description:**
 Fitness estimates (height, number of stems, flowers, fruits, fruit to
 flower ratio, herbivory, seed viability) of individual plants in common
 garden. This file contains 19 columns and 367 rows (obs) of data. #####
 Variables * SiteID: site name; geographic location of maternal plant
 accession (Text) Illinois Beach 2, Newark Road, Pine Station, Shaw
 Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * ColorS:
 dominant bract color of site of maternal plant acession grouped into two
 categories (Yellow or Orange) * PlantID: maternal plant unique numerical
 ID * Treatment: type of observation, (text) recorded as either Morphology
 or Fitness * ColorM(RHS): color of materal plant, alpha numeric Royal
 Horticulture Society code for color * ColorM: color of maternal plant,
 plant bract color grouped into two categories (Yellow or Orange) *
 CommGDID: individual plant in common garden unique numerical ID *
 ColorP(RHS): color of plant, alpha numeric Royal Horticulture Society code
 for color * ColorP: plant bract color grouped into two categories (Yellow
 or Orange) * Height: height of plant (cm) * Stems: number of stems
 (integer) * Flowers: number of flowers (integer) * Fruit: number of fruits
 (integer) * Herbivory: categorical estiamte of herbivory, values = TRUE,
 FALSE * Failures: number of flowers that failed to set fruit * Ratio:
 ratio of fruits to flowers * ColorSM: variables Color S and ColorM
 combined * ColorSP: vairables ColorS and ColorP combined * ColorSMP:
 variables ColorS, ColorM and ColorP combined #### File:
 Fruit_flower_Field_Herbivory.csv **Description:** Herbivory estimates of
 individual plants at field sites. This file contains 13 columns and 152
 rows (obs) of data. ##### Variables * Site: site name; geographic location
 of observation (Text) Illinois Beach 2, Newark Road, Pine Station, Shaw
 Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * ColorS:
 dominant bract color of site grouped into two categories (Yellow or
 Orange) * PlantID: unique numerical ID for each individual plant *
 Treatment: type of observation, (text) recorded as either Morphology,
 Fitness or Herbivory * Color(RHS): alpha numeric Royal Horticulture
 Society code for color * ColorP: plant bract color grouped into two
 categories (Yellow or Orange) * LeavesNo: number of leaves (integer) *
 DamageL: number of leaves damaged by herbivory (integer) * FlowersNo:
 number of flowers (integer) * DamageFl: number of flowers damaged by
 herbivory (integer) * ColorSP: combination of Site color (ColorS =
 dominant bract color of site grouped into two categories (Yellow or
 Orange)) and plant color (ColorPwith individual plant bract color grouped
 into two categories (Yellow or Orange)) * PropLeaves: proportion of
 damaged leaves to total leaves (ratio) * PropFlowers: proportion of
 damaged flowers to total flowers (ratio) #### File:
 Fruit_flower_Field_Seed.csv **Description:** Seed counts (capsule fill)
 estimates of individual plants at field sites. This file contains 11
 columns and 176 rows (obs) of data. ##### Variables * Population: site
 name (Text); geographic location of observation Illinois Beach 2, Newark
 Road, Pine Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie,
 Hoosier Prairie, Meissner-Corron, Newark Road, Sand Ridge,
 Gensburg-Markham * ColorS: dominant bract color of site grouped into two
 categories (Yellow or Orange) * Plant ID #: unique numerical ID for each
 individual plant * Treatment: type of observation, (text) recorded as
 either Morphology or Fitness * Bract Color: alpha numeric Royal
 Horticulture Society code for color * ColorP: plant bract color grouped
 into two categories (Yellow or Orange) * Total: total number of seed
 capsules * AvgOfFilled (&gt;50%): average number of seed capsules that
 were more than 50% filled * AvgOfEmpty (&lt;50%): average number of seed
 capsules that were less than 50% filled * ratio: ratio of filled to total
 seed capsules * ColorSP: combination of site color (ColorS = dominant
 bract color of site grouped into two categories (Yellow or Orange)) and
 plant color (ColorP = individual plant bract color grouped into two
 categories (Yellow or Orange)) #### File:
 CACO_JF_Colorofplants_20241111.csv **Description:** Index of data
 collection for morphology, fitness, or herbivory by individual plant ID
 and site. This file contains 7 columns and 541 rows (obs) of data. #####
 Variables * Plant ID: unique numerical ID for each individual plant *
 SiteID : site name; geographic location of observation (Text) Illinois
 Beach 2, Newark Road, Pine Station, Shaw Prairie, llinois Beach 1,
 Dropseed Prairie, Hoosier Prairie, Meissner-Corron, Newark Road, Sand
 Ridge, Gensburg-Markham * Year: year of data collection * Date: date of
 data collection * Treatment: type of observation, (text) recorded as
 either Morphology or Fitness or Herbivory * Broad Color ROY: plant bract
 color grouped into three categories (Red or Orange or Yellow) * Bract
 Color: alpha numeric Royal Horticulture Society code for color #### File:
 CACO_JFdata_NMDScg_20241023.csv **Description:** Floral morphology data
 for common garden plants (input for NMDS). This file contains 19 columns
 and 841 rows of data. ##### Variables * Site Name: site name; geographic
 location of observation (Text) Illinois Beach 2, Newark Road, Pine
 Station, Shaw Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * Plant ID:
 unique numerical ID for maternal plant * Maternal Bract Color: alpha
 numeric Royal Horticulture Society code for maternal bract color * Common
 Garden ID: umerical ID for common garden * RHS Code: alpha numeric Royal
 Horticulture Society code for color * ID1: unique numerical ID for each
 individual plant * CGID: umerical ID for common garden * Bract Length:
 length of bract from bact base to lobe tips * Bract Base: length of bract
 base to lobe base * Bract Width: width of bract at widest pointt * Bract
 Lobe: count of number of bract lobes * Calyx Outer: length of outer calyx
 segment (mm) * Calyx Inner: length of inner calyx segment (mm) * Corolla
 Tube Length: length of corrolla tube (mm) * Corolla Galea Length: length
 of corrolla galea (mm) * Corolla Tube Width: width of corrolla tube (mm) *
 Stamen Length: stamen length (mm) * Style Length: style length (mm) *
 Herkogamy: separation of anthers and stigma (stamen length - stigma
 length)) #### File: CACO_JFdata_NMDSfield_20241023.csv **Description:**
 Floral morphology data for field plants (input for NMDS). This file
 contains 18 columns and 532 rows (obs) of data. ##### Variables * Site
 Name: site name; geographic location of observation (Text) Illinois Beach
 2, Newark Road, Pine Station, Shaw Prairie, llinois Beach 1, Dropseed
 Prairie, Hoosier Prairie, Meissner-Corron, Newark Road, Sand Ridge,
 Gensburg-Markham * Plant ID: unique numerical ID for each individual plant
 with decimal indicating the flower measured on each plant * Bract Color:
 alpha numeric Royal Horticulture Society code for color * ID: unique
 numerical ID for each individual plant * Plant ID: unique numerical ID for
 each individual plant * Tag No: unique numerical ID for each individual
 plant * Bract Length: length of bract from bact base to lobe tips * Bract
 Base: length of bract base to lobe base * Bract Width: Witch of bract at
 widest pointt * Bract Lobe: count of number of bract lobes * Calyx Outer:
 length of outer calyx segment (mm) * Calyx Inner: length of inner calyx
 segment (mm) * Corolla Tube Length: length of corrolla tube (mm) * Corolla
 Galea Length: length of corrolla galea (mm) * Corolla Tube Width: width of
 corrolla tube (mm) * Stamen Length: stamen length (mm) * Style Length:
 style length (mm) * Herkogamy: separation of anthers and stigma (stamen
 length - stigma length)) #### File: Fruit_flower_Field.csv
 **Description:** Fruit-to-flower ratio of individual plants at field sites
 This file contains description 12 columns and 178 rows of data. #####
 Variables * rows (obs): 178 * SiteID: site name; geographic location of
 observation (Text) Illinois Beach 2, Newark Road, Pine Station, Shaw
 Prairie, llinois Beach 1, Dropseed Prairie, Hoosier Prairie,
 Meissner-Corron, Newark Road, Sand Ridge, Gensburg-Markham * ColorS:
 dominant bract color of site grouped into two categories (Yellow or
 Orange) * PlantID: unique numerical ID for each individual plant *
 Treatment: type of observation, (text) recorded as either Morphology or
 Fitness * ColorP(RHS): alpha numeric Royal Horticulture Society code for
 color * ColorP: plant bract color grouped into two categories (Yellow or
 Orange) * Height: height of flowering stem (cm) * Flowers: number of
 flowers (integer) * Fruit: number of fruits (integer) * Failures: number
 of flowers that failed to set fruit (integer) * Ratio: ratio of number of
 fruits to number of flowers for each plant * ColorSP: combination of Site
 color (ColorS = dominant bract color of site grouped into two categories
 (Yellow or Orange)) and plant color (ColorPwith individual plant bract
 color grouped into two categories (Yellow or Orange)) #### File:
 ColorChart.csv **Description:** Color conversion chart for RGB, HSV. CIE,
 hex codes and UCL colors of individual plants in the field. This file
 contains 23 columns and 236 rows (obs) of data. ##### Variables * ColorID
 : Unique identification number for each observation * RHS.: Royal
 Horticultural Society (RHS) color page number and letter * RHS.page.:
 Royal Horticultural Society (RHS) color page number * RHS.Hue: Royal
 Horticultural Society (RHS) letter * Color.Vector.distance: Bract color
 RGB values were transformed into a single vector value by treating each
 color as a point in n-dimensional space and determining the Euclidean
 distance of each point from the origin (zero). This was calculated by
 summing the squares of the three individual RGB values and calculating the
 square root of the total. * Field1: Field assesment of color, categorical
 as Orange, Red or Yellow * Color : Field assesment of color, categorical
 as Yellow, Greenish Yellow, Orangish Yellow, Yellowish pink, Orange,
 Yellowish Green, Reddish Orange, Yellowish Pink, Pink, Red, Purplish Red,
 Purplish Pink * UCL.: Universal Color Language * UCL.name: Universal Color
 Language name * R: Red (RGB) colorspace * G: Green (RGB) colorspace * B :
 Blue (RGB) colorspace * hex.RGB: Hexcode from RGB colorspace * R..sRGB :
 sRGB (standard RGB) colorspace R value (red value) * G..sRGB: sRGB
 (standard RGB) colorspace G value (green value) * B.sRGB: sRGB (standard
 RGB) colorspace B value (blue value) * L...CIE.Lab: CIELAB color space L
 value or lightness value * a...CIE.Lab: CIELAB color space a value or
 green value * b...CIE.Lab: CIELAB color space b value or blue value *
 L....CIE.LCh: CIE LCh Colour Space L value or lightness value *
 C....CIE.LCh : CIE LCh Colour Space C value * h....CIE.LCh: CIE LCh Colour
 Space h value #### File: KimDiss_CACOcross20250114b.csv **Description:**
 Results from controlled crosses in the common garden. This file contains 8
 columns and 17 rows (obs) of data. ##### Variables * MaternalColCatBroad
 maternal plant color as as "Red" or Yellow * MaternalColCatFine
 maternal plant color "Scarlet" or "Orange" or
 "Yellow" * Pollen_donor pollen donor color as "Red" or
 Yellow * Pollen_donor_fine pollon donor color "Scarlet" or
 "Orange" or "Yellow" * OffspringColor offspring color
 as "Red" or Yellow * OffspringColorFine offspring color as
 "Scarlet" or "Orange" or "Yellow" * count
 count of offspring * n_crosses count of crosses ### Habitat Data *All
 files contained in the Habitat_data.zip directory* #### File: Soil.csv
 **Description:** Herbarium soil and habitat properties. This file contains
 18 columns and 427 rows (obs) rows of data. ##### Variables * ID: Unique
 identifier for each plant * GPS.North: GPS coordinates (latitude) *
 GPS.West: GPS coordinates (longitude) * County: County where the sample is
 located (character) * State: State where the sample is located (character)
 * Confirmed: Confirmation status of bract color (TRUE or FALSE) * ColorS:
 Dominant bract color of site of maternal plant acession grouped into two
 categories (Yellow or Orange) * Texture_short: Simplified soil texture
 classification (factor; loam, sandy clay, silt loam, sandy loam, silt clay
 loam) * Eco.I: EPA Eco-reigon 1 classification (integer; 5, 8 or 9) *
 Saturation: Soil saturation level (integer) * Field_Capacity: Field
 capacity of the soil (integer) * Available_water: Amount of water
 available in the soil (integer) * Texture_broad: Broad soil texture
 classification (factor; clay loam muck, sand, other) * Wet.Area: Indicates
 if the area is is described as wet or wetland on herabrium label (TRUE or
 FALSE) * DRY: Indicates if the area is is described as dry on herabrium
 label (TRUE or FALSE) * Combined: Combined classification of soil (TRUE or
 FALSE) * Forested: Indicates if the area is described as forested on
 herabrium label (TRUE or FALSE) * Prairie: Indicates if the area is is
 described as prairie on herabrium label (TRUE or FALSE) #### File:
 Popaverages.csv **Description:** Field site characteristics (soil
 moisture, texture, cover class and plant communities). This file contains
 columns and rows (obs) of data. ##### Variables * Site: site name * Code:
 abreviated site name relating to geographic location of observation IB2 =
 Illinois Beach 2, MW = Newark Road, = Pine Station, SP = Shaw Prairie, IB1
 = Illinois Beach 1, DP = Dropseed Prairie, HP = Hoosier Prairie, MC =
 Meissner-Corron, NR = Newark Road, SR = Sand Ridge, GM = Gensburg-Markham
 * Site#: unique numerical identifier of site * GPS North : north–south
 position of a point * GPS West: east–west position of a point * Color S:
 dominant bract color grouped into two major categories Red or Yellow *
 Color Sx: dominant bract color grouped into three categories Red or Yellow
 or Polymorphic * County: describes geographic location of observation *
 State: describes geographic location of observation * Percent Yellow:
 percent of individuals recorded as yellow * Yellow: number of individuals
 recorded as having yellow bracts * Red: number of individuals recorded as
 having yellow bracts * BareSoil: proportion of ground in quadrat that was
 classfied as having no vegetation * FineDebris: proportion of ground in
 quadrat that was classfied as having fine debris * Dry (UPL or FACU):
 proportion of plants identified in community surveys as being obligate or
 facultatice upland plants * Wet (OBL or FACW): proportion of plants
 identified in community surveys as being obligate or facultative wetland
 plants * NRCS Soil: soil category as defined by National Resource
 Conservation Service * Soil Moisture Content: percent soil volumetric
 water content (VWC) * % Sand: percent of sand in soil * % Silt: percent of
 silt in soil * % Clay: percent of clay in soil #### File: Phenology.csv
 **Description:** Phenology of herbarium observations. This file contains 8
 columns and 2854 rows (obs) of data. ##### Variables * ID: Unique
 identifier for each observation * Accession.ID: Accession or catalog
 identifier for the sample * GPS.North: GPS coordinates (latitude) *
 GPS.West: GPS coordinates (longitude) * State: State of collection
 (character) * Julian.Days: Day of the year (Julian date) * ColorS:
 Dominant bract color of site of maternal plant acession grouped into two
 categories (Yellow or Orange) * Confirmed: Confirmation status of bract
 color (TRUE or FALSE) #### File: HabitatII.csv **Description:** Site
 characteristics from herbarium labels and EPA Ecoregions This file
 contains 14 columns and 1343 rows (obs) of data. ##### Variables * ID:
 Unique identifier for each observation * GPS.North: GPS coordinates
 (latitude) * GPS.West: GPS coordinates (longitude) * County: County where
 the sample is located * State: State where the sample is located * Eco.I:
 EPA Eco-reigon 1 classification (integer; 5, 8 or 9) * ColorS: Dominant
 bract color of site grouped into two categories (Yellow or Orange) *
 Confirmed: Indicates confirmation status of bract color (TRUE or FALSE) *
 Wet.Area: Indicates if the area is is described as wet or wetland on
 herabrium label (TRUE or FALSE) * DRY: Indicates if the area is is
 described as dry on herabrium label (TRUE or FALSE) * Combined: Combined
 classification of soil (TRUE or FALSE) * Forested: Indicates if the area
 is described as forested on herabrium label (TRUE or FALSE) * Prairie:
 Indicates if the area is is described as prairie on herabrium label (TRUE
 or FALSE) * Label.info: Additional information about the sample or label
 #### File: Nutrients.csv **Description:** Soil nutrient supply rates
 measured using Plant Root Simulator (PRS®) probes at multiple *Castilleja
 coccinea* field sites.\ This file contains 17 columns and 12 rows
 (observations) of data. ##### Variables * Site: Site code abbreviation
 where the PRS® probes were deployed * Color: Predominant bract color morph
 at the site (Red or Yellow) * NO3-N: Nitrate nitrogen (µg per 10 cm² per
 burial length) * NH4-N: Ammonium nitrogen (µg per 10 cm² per burial
 length) * Ca: Calcium (µg per 10 cm² per burial length) * Mg: Magnesium
 (µg per 10 cm² per burial length) * K: Potassium (µg per 10 cm² per burial
 length) * P: Phosphorus (µg per 10 cm² per burial length) * Fe: Iron (µg
 per 10 cm² per burial length) * Mn: Manganese (µg per 10 cm² per burial
 length) * Cu: Copper (µg per 10 cm² per burial length) * Zn: Zinc (µg per
 10 cm² per burial length) * B: Boron (µg per 10 cm² per burial length) *
 S: Sulfur (µg per 10 cm² per burial length) * Pb: Lead (µg per 10 cm² per
 burial length) * Al: Aluminum (µg per 10 cm² per burial length) * Cd:
 Cadmium (µg per 10 cm² per burial length) ##### Notes * Units represent
 nutrient supply rates as accumulated on PRS® probes during the burial
 period. * Zeros indicate values below detection limits for the assay. *
 Site codes (e.g., DP, GM, IB1) uniquely identify sampling locations. *
 Bract color (Red or Yellow) refers to the predominant floral morph
 observed at each site. ### Population genomics data *All files contained
 in the Genetics_denovo_data.zip directory* #### File:
 populations.snps_denovo_ustacks_R80_all.vcf **Description:** Unfiltered
 VCF file from STACKS de_novo run with r = 80 #### File:
 populations.snps_R80_ustacks_filt_filt_85miss.vcf.gz **Description:**
 Filtered VCF file contain SNPS for 37 individuals across 10 populations
 filtered based on these parameters: r = 80 (STACKS populations), Min depth
 of 5, Min genotype quality of 20, Allele balance between 0.25 and 0.75,
 Max depth of 50, Missing by sample cutoff 90%, Missing by SNP 85%, Min MAC
 3 #### File: admixture_input_051525.fam **Description:** Input for
 ADMIXTURE program run. Filtered for linkage disequilibrium with PLINK2.
 #### File: CACO_admix_K2_rep20_thin.Q **Description:** Output of ADMIXTURE
 program run for K = 2. #### File: mapmixture_gps_EM.csv **Description:**
 GPS coordinates of sites used to create admixture map (Figure 2) This file
 contains 3 columns and 10 rows rows of data. ##### Variables * Site:
 abbreviated site name relating to geographic location of observation IB2 =
 Illinois Beach 2, MW = Newark Road, = Pine Station, SP = Shaw Prairie, IB1
 = Illinois Beach 1, DP = Dropseed Prairie, HP = Hoosier Prairie, MC =
 Meissner-Corron, NR = Newark Road, SR = Sand Ridge, GM = Gensburg-Markham
 * lat: Latitude of the site * long: Longitude of the site #### File:
 mapmixture.csv **Description:** Genetic cluster assignments from the
 CACO_admix_K2_rep20_thin.Q file in the mapmixture format. This file
 contains 37 columns and 4 rows of data. ##### Variables * Site: population
 assignment * Individual: unique alphanumeric code for each individual in a
 population * Pop1: proportion of ancestry assignment in cluster 1 * Pop2:
 proportion of ancestry assignment in cluster 2 ## Access information Data
 was derived from the following sources: * Raw sequenced data from the
 ddRADseq study are archived at the NCBI Sequence Read Archive (SRA) under
 BioProject ID PRJNA1216984
 ([http://www.ncbi.nlm.nih.gov/bioproject/1216984](http://www.ncbi.nlm.nih.gov/bioproject/1216984)). * iNaturalist observation data downloaded via Global Biodiversity Information Facility (GBIF): * Research grade iNaturalist observations of *Castilleja coccinea* (*n = 2757;* iNaturalist contributors, 2024) were downloaded from the Global Biodiversity Information Facility (GBIF; [https://www.gbif.org/](https://www.gbif.org/)) in February of 2024 (doi.org/10.15468/dl.bnunfa) for processing with the ColorSelector pipeline (Luong et al., 2023). References GBIF.org. 2024. Global Biodiversity Information Facility Occurrence Download. Available at [https://doi.org/10.15468/dl.bnunfa](https://doi.org/10.15468/dl.bnunfa) [accessed 08 February 2024]. iNaturalist contributors, iNaturalist (2024). iNaturalist Research-grade Observations. iNaturalist.org. Occurrence dataset [https://doi.org/10.15468/ab3s5x](https://doi.org/10.15468/ab3s5x) accessed via GBIF.org on 2024-02-08. Luong, Y., A. Gasca-Herrera, T. M. Misiewicz, and B. E. Carter. 2023. A pipeline for the rapid collection of color data from photographs. *Applications in Plant Sciences* 11: e11546