DOI: 10.5061/dryad.hqbzkh1r4
title: Intraspecific phytochemical diversity increases with productivity but has mixed effects on herbivory

description: Resources play an important role in shaping the evolution of plant
 defensive strategies. How resource availability influences overall
 phytochemical diversity within species, and how this in-turn affects
 herbivore damage is not well understood. Using Monarda fistulosa, a
 species that produces diverse terpene compounds and distinct chemotypes
 (i.e., thymol or carvacrol), we addressed three questions: 1) How do
 phytochemical components (i.e., total concentration, Shannon diversity,
 and chemotypes) vary between distinct populations from two large, widely
 separated regions that differ in climate and productivity? 2) How do these
 components influence attack by different herbivore species? 3) What are
 the growth costs of higher phytochemical levels in different chemotypes?
 Seeds were collected from 12 populations spread across two regions that
 differ in precipitation-driven productivity, six from a low-productivity
 environment in Montana and six from a high-productivity environment in
 Wisconsin, and grown in a common garden in Wisconsin. We analyzed terpene
 concentrations, quantified the number of herbivores and damage, and
 measured plant above-ground biomass. Plants from Montana exhibited higher
 terpene concentrations but lower diversity, whereas those from Wisconsin
 displayed lower terpene concentrations but higher diversity. Plants from
 Wisconsin received more damage from a specialist seed predator and
 leaf-chewing herbivores, although abundance of a specialist aphid was
 lower, compared to plants from Montana. Total terpene concentration
 emerged as the primary predictor of herbivore damage, although some
 differences were observed between region of origin and with terpene
 richness. Herbivores exhibited mixed responses to phytochemistry; some
 herbivores were negatively affected, while a specialized leaf galler
 responded positively. Costs of producing defenses were evident by negative
 correlations between plant biomass and total terpene concentration (but
 not chemical richness or Shannon diversity). Our study revealed that
 regional differences in productivity can influence the evolution of
 phytochemical concentration and diversity, which, in turn, affects plant
 damage and incurs growth-related costs.
description: Data was collected in a common garden and laboratory. Data files
 are raw data and processed with the associated R script. Detailed methods
 are described in the associated paper.
description: # Intraspecific phytochemical diversity increases with resources
 availability but has mixed effects on herbivory Data for Monarda fistulosa
 phytochemistry and herbivory common garden project. The common garden
 experiment was located in Wisconsin, USA using seeds collected from M.
 fistulosa populations originating from Montana, USA and Wisconsin, USA.
 Chemistry was conducted on leaves to measure terpenes and herbivory was
 measured on the same plants. Contact Phil Hahn
 ([hahnp@ufl.edu](mailto:hahnp@ufl.edu)) with any questions. This
 manuscript has been accepted for publication (10/21/2024): Calixto, E.S.,
 J.L. Maron, K. Keefover-Ring, J.H. Cammarano, and P.G. Hahn. Intraspecific
 phytochemical diversity increases with resources availability but has
 mixed effects on herbivory. Oikos (accepted). 10.1111/oik.10914 R
 scripts (file: Monarda_ChemDivDmg_2020907.R) associated with these data
 files can be found here:
 [https://doi.org/10.5281/zenodo.13988961](https://doi.org/10.5281/zenodo.13988961) Data files include: ## Monarda\_Garden\_Traits\_2019.csv This file contains data on Monarda fistulosa leaf traits and phytochemistry. All data were collected by the authors. Blank cells represent missing data. | variables | units | description | | :------------------ | :--------------------: | :---------------------------------------------------------------------------------------------------- | | Region | categorical | Geographical region, MT = Montana, WI = Wisconsin | | Site | categorical | Code for site location, see file 'WIMT\_Sites\_2019.csv' or Table S1 for more details about locations | | Plant | number | numerical code for each maternal plant for where seed was collected | | Rep | categorical | A-E, code for indivdual plants grown from the maternal plant ID (ie. half siblings) | | Row | number | row in common garden, used to locate plants in the garden | | OrderGard | number | order planted in garden, used to locate plants in the garden | | Unique | number | unique number for each plant | | Biomass\_g | g | dry biomass of plants harvested on 13 August 2019 | | codedup | categorical | Plant and Rep column pasted together | | Chemo | categorical | chemotype, T = thymol, C = carvacrol | | a\_thujene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | a\_pinene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | sabinene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | b\_pinene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | octen\_3\_ol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | myrcene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | a\_phellandrene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | carene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | a\_terpinene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | p\_cymene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | limonene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | beta\_ocimene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | gama\_terpinene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | cis\_sab\_hydrate | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | trans\_sab\_hydrate | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | borneol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | terp\_4\_ol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | cym\_8\_ol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | a\_terpineol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | nerol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | isoborneol\_formate | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | thymoquinone | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | geraniol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | geranial | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | thymol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | carvacrol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | eugenol | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | caryophyllene | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | germ\_D | mg / g dry leaf tissue | concentration of chemical compound, see Table S2 for full compound name additional info | | total | mg / g dry leaf tissue | concentration of total chemical compounds | | totTC | mg / g dry leaf tissue | concentration of only thymol and carvacrol | ## MonardaGarden\_Data2021.csv This file contains data on Monarda plant measurements in the field. All data were collected by the authors. Blank cells represent missing data. | variables | units | description | | :---------- | :---------: | :---------------------------------------------------------------------------------------------------- | | Region | categorical | Geographical region, MT = Montana, WI = Wisconsin | | Site | categorical | Code for site location, see file 'WIMT\_Sites\_2019.csv' or Table S1 for more details about locations | | Plant | number | Identifier for plant sample | | Rep | number | Replication number | | Row | number | Row position in the field | | Order | number | order planted in garden, used to locate plants in the garden | | hg\_Tallest | cm | Height of the tallest plant part | | volD1 | cm | Diameter of plant at widest point, used to estimate volume | | volD2 | cm | Diameter of plant orthogonal to volD1, used to estimate volume | | b | presence | Presence (1) or absence (0) of buds | | fl | presence | Presence (1) or absence (0) of flowers | | fr | presence | Presence (1) or absence (0) of fruits | | infl | count | Number of inflorescences | | ch | % | Percent of chewing damage estimated on whole plant | | prc | % | Percent of piercing/sucking damage estimated on whole plant | | notes | | Additional notes or observations | ## MonardaGarden\_SeedDamage2021 (5).csv This file contains data on seed head damage to Monarda plants in the common garden. All data were collected by the authors. Blank cells represent missing data. | variables | units | description | | :-------- | :---------: | :---------------------------------------------------------------------------------------------------- | | Region | categorical | Geographical region, MT = Montana, WI = Wisconsin | | Site | categorical | Code for site location, see file 'WIMT\_Sites\_2019.csv' or Table S1 for more details about locations | | Plant | number | Identifier for plant sample | | Rep | number | Replication number | | Row | number | Row position in the field | | Flowered | Presence | Whether the plant flowered | | HeadSize | cm | Size of the flower head | | HeadMass | g | Mass of the flower head | | N\_tubes | count | Number of tubes | | HerbDmg | % | Percentage of herbivory damage | | Col\_grub | count | Number of Coleoptera grubs | | Lep\_grub | count | Number of Lepidoptera grubs | | Lep\_Phy | count | Number of Phytophagous Lepidoptera | | Fly | count | Number of flies | | Unknwn | count | Number of unknown organisms | | Seeds\_No | count | Number of seeds | | Seed\_wt | g | Weight of the seeds | | Notes | | Additional notes or observations | | DateP | Date | Date of the recorded measurement | ## MonardaGarden2020.csv This file contains data collected in the common garden site in Wisconsin in the summer of 2020. All data were collected by the authors. Blank cells represent missing data. | variables | units | description | | :-------------------- | :--------------: | :---------------------------------------------------------------------------------------------------- | | Region | categorical | MT = Montana, WI = Wisconsin | | Site | categorical | Code for site location, see file 'WIMT\_Sites\_2019.csv' or Table S1 for more details about locations | | Plant | number | numerical code for each maternal plant for where seed was collected | | Rep | categorical | A-E, code for indivdual plants grown from the maternal plant ID (ie. half siblings) | | Row | number | row in common garden, used to locate plants in the garden | | Order | number | order planted in garden, used to locate plants in the garden | | survey | category | month of survey, July or Aug=August | | height | cm | stem height, ground to top of plant | | leaves\_total | count | number of leaves on plant | | leaves\_damaged | count | number of leaves with damage | | herbpercentage | percent | overall estimate of percent damage to the whole plant | | leaf\_1 | percent | percent damage visually estimated on a single leaf | | leaf\_2 | percent | percent damage visually estimated on a single leaf | | leaf\_3 | percent | percent damage visually estimated on a single leaf | | leaf\_4 | percent | percent damage visually estimated on a single leaf | | leaf\_5 | percent | percent damage visually estimated on a single leaf | | leaf\_6 | percent | percent damage visually estimated on a single leaf | | leaf\_7 | percent | percent damage visually estimated on a single leaf | | leaf\_8 | percent | percent damage visually estimated on a single leaf | | leaf\_9 | percent | percent damage visually estimated on a single leaf | | leaf\_10 | percent | percent damage visually estimated on a single leaf | | mildew | presence absence | presence of powdery mildew, 0=absent, 1=present | | mines\_blotch | count | number of powdery mine blotches | | mines\_linear | count | number of leaf mines | | galls\_leaf | count | number of leaf galls by Ametrodiplosis fistulosae | | galls\_stem | count | number of stem galls | | galls\_bud | count | number of bud galls | | leps | count | number of lepidoptera caterpillars counted on plant | | aphids\_parasitized | count | number of parasitized aphid mummies on plant | | aphids\_unparasitized | count | number of live aphids (Aphis monardae) on plant | | ants | count | number of ants on plant | | ant\_genus | category | genus of ant | | flowerbuds | count | number of flower buds | | bloom\_date | date | date of bloom | | damage\_chewing | percent | percent chewing damage to whole plant | | damage\_spot | presence | presence of spotting damage | | agromyzid | count | number of agromyzid flies on plant | ## WIMT\_Sites\_2019.csv This file contains information on site characteristics for locations where seed was originally collected. Soil data were obtain from: 10.5061/dryad.x0k6djhjq. Variables indicated as bioclim variables were obtained from: [https://www.worldclim.org/data/bioclim.html](https://www.worldclim.org/data/bioclim.html). Blank cells represent missing data. | variables | units | description | | :-------- | :---------: | :--------------------------------------------------- | | Region | categorical | Geographical region, MT = Montana, WI = Wisconsin | | Site | categorical | Code for site location | | Lat | degrees | Latitude coordinate | | Long | degrees | Longitude coordinate | | pH | | Soil pH level | | orgmat | % | Organic matter content | | nit | % | Nitrogen content | | K | % | Potassium content | | cec | meq/100g | Cation exchange capacity | | P | % | Phosphorus content | | mat1 | degrees C | bioclim1, mean annual temperature | | iso3 | | bioclim3, isothermality | | tsd4 | | bioclim4, standard deviation of monthly temperatures | | map12 | cm/year | bioclim12, Mean annual precipitation | | pwrq18 | | bioclim18, precipication in the warmest quarter |