DOI: 10.5061/dryad.jm63xsjnp
title: Data from: Persistence of the ecological niche in pond damselflies underlies a stable adaptive zone despite varying selection

description: Following the development of regression-based methods to estimate natural
 and sexual selection, evolutionary biologists have quantified the
 strength, mode, and direction of selection in natural populations.
 Although this approach has been successful, its limitations include the
 lack of replication across species, which compromises the generality of
 the inferences beyond microevolutionary time scales. Here, we carried out
 a comparative study of selection on wing shape and body size across
 multiple populations of two closely related pond damselflies: Enallagma
 cyathigerum and Ischnura elegans (Odonata: Coenagrionidae). We found weak
 stabilizing selection on wing shape in both sexes, and no evidence that
 selection on this trait differed between the species. In contrast, the
 selection of body size was curvilinear in males and directional in
 females, with differences in form (males) and intensity (females) between
 the species. By analyzing selection in each local pond, we found that
 selection on male body size was shaped by various population
 characteristics that reflect local mating systems. Furthermore, the
 relationship between mating system characteristics and directional
 selection was remarkably consistent across these species. Finally, we
 present a graphical model that links contemporary selection with
 macroevolution. Based on this model, we conclude that the persistence of
 ecological modes of life in pond damselflies explains why varying
 selection in nature may still result in a stable adaptive zone lasting
 millions of years.
description: # Persistence of the ecological niche in pond damselflies underlies a
 stable adaptive zone despite varying selection **Authors** Anjali Gupta,
 Erik I. Svensson, Henrik Frietsch and Masahito Tsuboi **Contact**
 [masa.tsuboi@gmail.com](mailto:masa.tsuboi@gmail.com) ### Description This
 data package contains morphological and ecological data of two pond
 damselfly species, *Ischnura elegans* and *Enallagma cyathigerum*, from
 southern Sweden. Data were obtained by the authors. Code was written by
 Anjali Gupta and Masahito Tsuboi. **Data Files** **Data is available in**
 1. **Is_updated.csv**: Data file for opportunity of sexual selection and
 male mating success - derived from community sampling from 2018-2021.
 Units are specified in the main text. A description to each column in the
 data can be found below - Species: *Ischnura elegans* / *Enallagma
 cyathigerum,* Locale: Location site of specimen collected,
 MaleMatingSuccess: mean proportion of male (belonging to one species)
 mating at a particular locale in one field season Is: opportunity for
 sexual selection ( variance in male mating success / ((average male mating
 success)^2) ) Year: Year/Field season of specimen collection n: frequency
 2. **DataPC_sorted_locale_fixed.csv**: Data file for wing shape and body
 size PC components with intuitive locale names to evaluate selection
 gradients. Units are specified in the main text. A description to each
 column in the data can be found below - Ind_ID: Unique ID for each
 specimen, Season_ID: ID for each specimen for a particular field season,
 Genus, Genus_, Species: Genus and species name for the specimen, Date:
 Date of specimen collection, Locale: Location site of specimen collected,
 Cop_ID: ID for individuals caught in a couple (the male and female caught
 in a couple get the same ID), NA for individuals that are caught not
 mating, Sex: Male/Female, Age: Mature/Immature based on age at the time of
 specimen collection, Copula_, Copula: Single (0)/ Couple (1) - based on
 whether the individual caught non-mating / mating, Thor_col: Thorax color,
 NA if not scored, Col_phase: NA for this dataset, Parasite: Number of
 mites on the body of the specimen when captured, Morph: Female color morph
 type, NA for males, Wing: Wing length of the specimen, Thorax: Thorax
 length of the specimen, Length: Total body length of the specimen,
 Abdomen: Abdomen length of the specimen, S_4: S4 segment / abdomenal width
 of the specimen, Eggs: No of eggs laid by a female caught in a couple (NA
 for males and single-caught females) Season: Year of field season
 WingSide: Right/left wing specimen WingID: Forewing/ Hindwing X0,Y0 -
 X16,Y16: X and Y coordinates of the 17 landmarks on wing GPA COORDS NEXT:
 NA for this data X.1,Y.1 - X.17,Y.17:  Generalized Procrustes analysis
 transformed X and Y coordinates of the 17 landmarks on wing 
 MatingSuccess: mean proportion of male (belonging to one species) mating
 at a particular locale in one field season Fecundity: mean-standardized
 eggs laid by a female, NA for males and single-caught females LD1:
 Component values of major axis - LD1 from linear discriminant function
 analysis on wing shape PC1_shape - PC5_shape: Component values of PC1-PC5
 axes from principal component analysis on wing shape PC1size -
 PC2_size: Component values of PC1-PC2 axes from principal component
 analysis on body size PC1_shape2 - PC5_shape2: Squared values of PC1_shape
 - PC5_shape PC1size2 - PC2_size2: Squared values of PC1size - PC2_size
 LD12: Squared values of LD1 3. **Data_LMRenamed.csv**: Data file for
 repeated measurements of same specimens to evaluate repeatability. Units
 are pixels. A description to each column in the data can be found below -
 Season_ID: ID for each specimen for a particular field season, Genus,
 Species: Genus and species name for the specimen, Date: Date of specimen
 collection, Locale: Location site of specimen collected, Cop_ID: ID for
 individuals caught in a couple (the male and female caught in a couple get
 the same ID), NA for individuals that are caught not mating, Sex:
 Male/Female, Age: Mature/Immature based on age at the time of specimen
 collection, Copula_, Copula: Single (0)/ Couple (1) - based on whether
 the individual caught non-mating / mating, Thor_col: Thorax color, NA if
 not scored, Col_phase: NA for this dataset, Parasite: Number of mites on
 the body of the specimen when captured, Morph: Female color morph type, NA
 for males, Wing: Wing length of the specimen, Thorax: Thorax length of the
 specimen, Length: Total body length of the specimen, Abdomen: Abdomen
 length of the specimen, S_4: S4 segment / abdomenal width of the specimen,
 Eggs: No of eggs laid by a female caught in a couple (NA for males and
 single-caught females) Season: Year of field season WingSide: Right/left
 wing specimen WingID: Forewing/ Hindwing X0,Y0 - X16,Y16: X and Y
 coordinates of the 17 landmarks on wing GPA COORDS NEXT: NA for this data
 X.1,Y.1 - X.17,Y.17:  Generalized Procrustes analysis transformed X and Y
 coordinates of the 17 landmarks on wing  4. **Data.csv**: Raw data for all
 specimens of Ischnura and Enallagma focused sampling. All data used for
 the presented analyses are in data file #2. This is for archiving purpose
 only. A description to each column in the data can be found below -
 Season_ID: ID for each specimen for a particular field season, Genus,
 Species: Genus and species name for the specimen, Date: Date of specimen
 collection, Locale: Location site of specimen collected, Cop_ID: ID for
 individuals caught in a couple (the male and female caught in a couple get
 the same ID), NA for individuals that are caught not mating, Sex:
 Male/Female, Age: Mature/Immature based on age at the time of specimen
 collection, Copula_, Copula: Single (0)/ Couple (1) - based on whether
 the individual caught non-mating / mating, Thor_col: Thorax color, NA if
 not scored, Col_phase: NA for this dataset, Parasite: Number of mites on
 the body of the specimen when captured, Morph: Female color morph type, NA
 for males, Wing: Wing length of the specimen, Thorax: Thorax length of the
 specimen, Length: Total body length of the specimen, Abdomen: Abdomen
 length of the specimen, S_4: S4 segment / abdomenal width of the specimen,
 Eggs: No of eggs laid by a female caught in a couple (NA for males and
 single-caught females) Season: Year of field season WingSide: Right/left
 wing specimen WingID: Forewing/ Hindwing X0,Y0 - X16,Y16: X and Y
 coordinates of the 17 landmarks on wing GPA COORDS NEXT: NA for this data
 X.1,Y.1 - X.17,Y.17:  Generalized Procrustes analysis transformed X and Y
 coordinates of the 17 landmarks on wing 5. **CommunitySampling_2018.csv**:
 Raw data for Odonata community sampling for 2018. Each entry represents a
 observation. A description to each column in the data can be found below -
 Date: Date of specimen collection, Time: Time of specimen collection,
 Locale: Location site of specimen collected, Species: Genus and species
 name for the specimen, Age: Mature/Immature based on age at the time of
 specimen collection, Morph: Female color morph type, NA for males,
 Copulation.status: Single (0)/ Couple (1) - based on whether
 the individual caught non-mating / mating, Parasites: Number of mites on
 the body of the specimen when captured, Catching.time.minutes.: Time (in
 minutes) spent during specimen collection during one field visit,
 Year: Year of field season, Ischunra.elegans.morph: *I. elegans* female
 color morph type, NA for males and other species,
 Ischunra.elegans.phase: *I. elegans* female color phase type based on
 thorax color, NA for males and other species 6.
 **CommunitySampling_2019.csv**: Raw data for Odonata community sampling
 for 2019. Each entry represents a observation. A description to each
 column in the data can be found below - Date: Date of specimen collection,
 Time: Time of specimen collection, Locale: Location site of specimen
 collected, Species: Genus and species name for the specimen,
 Age: Mature/Immature based on age at the time of specimen collection,
 Morph: Female color morph type, NA for males, Copulation.status: Single
 (0)/ Couple (1) - based on whether the individual caught non-mating /
 mating, Parasites: Number of mites on the body of the specimen when
 captured, Catching.time.minutes.: Time (in minutes) spent during specimen
 collection during one field visit, Year: Year of field season,
 Ischunra.elegans.morph: *I. elegans* female color morph type, NA for males
 and other species, Ischunra.elegans.phase: *I. elegans* female color phase
 type based on thorax color, NA for males and other species 7.
 **CommunitySampling_2020.csv**: Raw data for Odonata community sampling
 for 2020. Each entry represents a observation. A description to each
 column in the data can be found below - Date: Date of specimen collection,
 Time: Time of specimen collection, Locale: Location site of specimen
 collected, Species: Genus and species name for the specimen,
 Age: Mature/Immature based on age at the time of specimen collection,
 Morph: Female color morph type, NA for males, Copulation.status: Single
 (0)/ Couple (1) - based on whether the individual caught non-mating /
 mating, Parasites: Number of mites on the body of the specimen when
 captured, Catching.time.minutes.: Time (in minutes) spent during specimen
 collection during one field visit, Year: Year of field season,
 Ischunra.elegans.morph: *I. elegans* female color morph type, NA for males
 and other species, Ischunra.elegans.phase: *I. elegans* female color phase
 type based on thorax color, NA for males and other species 8.
 **CommunitySampling_2021.csv**: Raw data for Odonata community sampling
 for 2021. Each entry represents a observation. A description to each
 column in the data can be found below - Date: Date of specimen collection,
 Time: Time of specimen collection, Locale: Location site of specimen
 collected, Species: Genus and species name for the specimen,
 Age: Mature/Immature based on age at the time of specimen collection,
 Morph: Female color morph type, NA for males, Copulation.status: Single
 (0)/ Couple (1) - based on whether the individual caught non-mating /
 mating, Parasites: Number of mites on the body of the specimen when
 captured, Catching.time.minutes.: Time (in minutes) spent during specimen
 collection during one field visit, Year: Year of field season,
 Ischunra.elegans.morph: *I. elegans* female color morph type, NA for males
 and other species, Ischunra.elegans.phase: *I. elegans* female color phase
 type based on thorax color, NA for males and other species
 **Code/Software** **R Code is available in** 1.
 **JEB_#1_Data_Preparation_FINAL.R**: Code for Generalized Procustes
 Analysis, Principal Component Analysis, and landmarking repeatability.
 This code reorganizes **Data.csv** to produce
 **DataPC_sorted_locale_fixed.csv** 2.
 **JEB_#2_Global_Comparisons_FINAL.R**: Code to evaluate selection per
 species and choose best models. 3.
 **JEB_#3_Population_Comparisons_FINAL.R**: Code to evaluate selection per
 population and choose best models. 4. **JEB_#4_prereq_OppSel_FINAL.R**:
 Code to calculate opportunity for sexual selection and male mating success
 estimates. This code reorganizes **CommunitySampling_2018.csv**,
 **CommunitySampling_2019.csv**, **CommunitySampling_2020.csv**, and
 **CommunitySampling_2021.csv** to produce **Is_updated.csv** 5.
 **JEB_#4_Selection_and_ecology_FINAL.R**: Code to summarize mating system
 parameters per population per year per species, Table S8, Table S9, Figure
 5 and Figure S8. 6. **JEB_#5_OtherFigures_FINAL.R**: Code for Figure 3,
 Figure S6, Figure S7, Figure S10 7. **Figure S2.R**: Code for Figure S2.
 **Wing Images** **Wing images used in this study are archived in three
 folders** 1. **Ischnura2020.zip**: Wing images from *Ischnura elegans*
 samples collected in 2020. 2. **Ischnura2021.zip**: Wing images from
 *Ischnura elegans* samples collected in 2021. 3. **Enallagma2021.zip**:
 Wing images from *Enallagma cyathigerum* samples collected in 2021. 4.
 **TrainingSetImages.zip**: Wing images used to train ml-morph for
 landmarking. **All filenames in the wing images folders follow our custom
 nomenclature "oi_IMG_XXX_id_XXX_wi_XX.jpg"** oi = original image
 IMG_XXX = This 'XXX' is not informative and was just used for
 our records while imaging wings id_XXX = This 'XXX' corresponds
 to the original ID of the specimen in the **Data.csv** datasheet wi_XX =
 This 'XX' can take four values -
 'rh','lh','rf','lf' corresponding
 to right/left hindwing/forewing