DOI: 10.5061/dryad.hhmgqnkt8
title: Microgeographic diversity does not drive macroevolutionary divergence in bite force of the Ibiza wall lizard, Podarcis pityusensis

description: We investigate evolutionary scaling in form-function-environment
 relationships of Podarcis pityusensis, the Ibiza wall lizard. A
 total of 307 lizards from 11 populations were surveyed for bite force and
 morphology in September/October 2022 and May 2023. These measurements were
 analysed with ecological data including diet, physical environment,
 population density, and levels of aggression for each location to
 investigate evolutionary drivers of form and function in this species.
 Analysis was done across five levels of evolutionary isolation to
 investigate whether microgeographic variation in this species contributes
 towards phenotypic divergence on more macroevolutionary scales. We find
 general incongruence in form-function-environment relationships across
 scales investigated, with a lack of phenotypic divergence at the largest
 scale obscuring high microgeographic variation. Further, the specific
 environmental drivers of phenotype vary on the level of isolation
 investigated and on sex, with female phenotypes apparently more strongly
 shaped by natural selection and male phenotypes most strongly shaped by a
 combination of natural and sexual selection.
description: # Microgeographic diversity does not drive macroevolutionary divergence in
 bite force of the Ibiza wall lizard, *Podarcis pityusensis* Dataset DOI:
 [10.5061/dryad.hhmgqnkt8](10.5061/dryad.hhmgqnkt8) ## Description of the
 data and file structure Phenotypic measurements and ecological data for 11
 populations of the Ibiza wall lizard, *Podarcis pityusensis*. Data is used
 to analyse form-function-environment relationships across scales of
 evolutionary isolation. ### Files and variables #### File:
 Supplementary_Data_Sheet_1.csv **Description:** Study sites with
 corresponding sample sizes, clade classification and ecological metrics.
 ##### Variables * Location: Name of the location in the Pityusic islands
 at which measurements were taken. Non-accented characters are used to ease
 use of files in R. * Female_sample_size: Number of females sampled for
 phenotypic measurement. * Male_sample_size: Number of males sampled for
 phenotypic measurement. * Islet: Specifying whether the location is on a
 small islet or a main island (Ibiza and Formentera) * Clade:
 Classification according to clade as specified in Rodríguez et al., (2013)
 * Percent_plant: Percentage plant food in the diet, measured using scats
 collected from each location, measured according to methods outlined in
 Pérez-Cembranos, León and Pérez-Mellado (2016). * Percent_hard: Percentage
 of the diet classified as 'hard' measured using scats collected
 from each location. Types of prey measured according to methods outlined
 in Pérez-Cembranos, León and Pérez-Mellado (2016), see Supplementary Data
 Sheet 5, then assigned 'hard' or 'soft' as described
 in Supplementary Data Sheet 6. * Percent_cannibalistic: Percentage
 *Podarcis* in the diet, measured using scats collected from each location.
 Types of prey measured according to methods outlined in Pérez-Cembranos,
 León and Pérez-Mellado (2016), see Supplementary Data Sheet 5. *
 Dietary_diversity: Hill q=2 value * Female_amputation_rate: proportion *
 Male_amputation_rate: proportion * Female_tail_autotomisation_rate:
 proportion * Male_tail_autotomisation_rate: proportion *
 Overall_amputation_rate: proportion, both sexes *
 Overall_tail_autotomisation_rate: proportion, both sexes *
 Population_density: measured as described in the text, see Supplementary
 Data Sheet 4 for further information. * Log_population_density: Log
 transformed values of population density * Islet_area_ha: in hectares,
 gained as described in the text * Islet_max_altitude_m: in metres, gained
 as described in the text * Biotic_index: calculated as described in the
 text, according to methods described in (Cheylan, 1992; Parlanti et al.,
 1988). #### File: Supplementary_Data_Sheet_2.csv
 **Description:** Phenotypic measurements of each individual taken in the
 field. ##### Variables * CLADE: Classification according to clade as
 specified in Rodríguez et al., (2013) * LOCATION: Name of the location in
 the Pityusic islands at which measurements were taken. Non-accented
 characters are used to ease use of files in R. * Lighthouse: Listed for
 Bleda Plana and Conillera, describing whether the population was from the
 lighthouse or the shoreline. All other locations listed as NA. * LIZARD:
 Lizard number in the field * SEX: Male (M) or female (F) * GAPE_ANGLE:
 Estimation of gape angle made as described in the text * MAX: Bite force,
 in Newtons * TAIL: Whether the tail had been autotomised (indicated by
 'Dropped') or was intact (indicated by 'Intact'). *
 WEIGHT: in grams * SVL: Snout-Vent-Length, in mm * PL: Pileus length, in
 mm * PW: Pileus width, in mm * HH: Head height, in mm * MOUTH: Mouth
 width, in mm * AMPUTATION: Whether digits or phalanges were missing
 (indicated by 'Yes') or intact (indicated by 'No). ####
 File: Supplementary_Data_Sheet_3.csv **Description:** Further data on the
 bite force trials performed by all lizards. Each lizard performed three
 bite force trials. ##### Variables * DATE: in DD.MM.YYYY format * CLADE:
 Classification according to clade as specified in Rodríguez et al., (2013)
 * LOCATION: Name of the location in the Pityusic islands at which
 measurements were taken. Non-accented characters are used to ease use of
 files in R. * Lighthouse: Listed for Bleda Plana and Conillera, describing
 whether the population was from the lighthouse or the shoreline. All other
 locations listed as NA. * LIZARD: Lizard number in the field * SEX: Male
 (M) or female (F) * BITE_1: Bite force in Newtons of the first bite trial
 * BITE_2: Bite force in Newtons of the second bite trial * BITE_3: Bite
 force in Newtons of the third bite trial * AVERAGE: Average bite force in
 Newtons across all three trials * MAX: Maximum bite force in Newtons
 across all three trials #### File: Supplementary_Data_Sheet_4.csv
 **Description:** Data used to calculate population density at each
 location, including precise location information and the year of
 measurement. ##### Variables * locality: Name of the location in the
 Pityusic islands at which measurements were taken. Non-accented characters
 are used to ease use of files in R. * zone: Specific location information
 * year * month * contacts: Number of lizards detected during the transect
 * length: Length of transect, in metres * density: Population density,
 calculated as described in the text * SE: Standard error * AIC: Akaike
 Information Criterion * g(x): Detectability value #### File:
 Supplementary_Data_Sheet_5.csv **Description:** All diet data used for
 each population, with corresponding year and month. Each row represents
 one fece. ##### Variables * month: Month that the fece was collected *
 year: Year that the fece was collected * locality: Location where the fece
 was collected. Non-accented characters are used to ease use of files in R.
 * % plant matter: Percentage plant matter in the scat, measured according
 to methods outlined in Pérez-Cembranos, León and Pérez-Mellado (2016). *
 All other columns: Prey grouping, as specified by column header. Number of
 each type of prey listed, measured according to methods outlined in
 Pérez-Cembranos, León and Pérez-Mellado (2016). #### File:
 Supplementary_Data_Sheet_6.csv **Description:**  Information on the
 classification of food into 'hard' and 'soft', listing
 the reference the classification was gained from. Food items missing an
 entry in the 'reference' column were classified according to our
 own knowledge. #### File: Podarcis_pityusensis_bite_force_RScript.R
 **Description:** R script used for the analysis. ## Code/software An R
 script is provided which corresponds to the 'data analysis'
 section of the manuscript. Analysis was performed on R version 4.3.2
 (2023-10-31 ucrt) -- "Eye Holes" Copyright (C) 2023 The R
 Foundation for Statistical Computing. R packages used are as follows:
 ggplot2, stats, car, hillR, simboot, MASS, jmv, Matrix, remotes, RRPP.
 Other data is in .csv format. ## Access information This Dryad dataset
 refers to methods described in the following sources: * Cheylan ,
 M.(1992). Synthèse biogéographique. In M. Delaugerre & M. Cheylan
 (Eds.), Atlasde répartition des batraciens et rep- tiles de corse (pp.
 105–120).Montpellier, Parc Naturel Régional de Corse/École Pratique des
 Hautes Études. * Parlanti , C., Lanza, B., Poggesi, M., & Sbordoni, V.
 (1988). Anfibi e rettili delle isole del Mediterraneo: Un testdellípotesi
 dell’equilibrio insulare. Bulletin of Ecology , 19:335–348. *
 Pérez-Cembranos , A., León, A., & Pérez-Mellado, V. (2016). Omnivory
 of an insular lizard: Sources of variation in the diet of Podarcis
 lilfordi (Squamata, Lacertidae). PLoS ONE , 11 , e0148947. https: //
 doi.org/ 10.1371/ journal.pone.0148947 This Dryad dataset includes data
 derived from the following sources: * Vanhooydonck, B., Herrel, A., Van
 Damme, R., Reilly, S.M., McBrayer, L.D., Miles, D.B., 2007. Interactions
 between habitat use, behavior, and the trophic niche of lacertid lizards.
 Lizard ecology: the evolutionary consequences of foraging mode 427–449. *
 Donihue, C.M., 2016. Aegean wall lizards switch foraging modes, diet, and
 morphology in a human-built environment. Ecology and Evolution 6,
 7433–7442. * Lluch i Dubon, 1997. Geografia de les Illes Balears. Lleonard
 Muntaner Editor. * Rodríguez, V., Brown, R.P., Terrasa, B., Pérez‐Mellado,
 V., Castro, J.A., Picornell, A., Ramon, M.M., 2013. Multilocus genetic
 diversity and historical biogeography of the endemic wall lizard from
 Ibiza and Formentera, *Podarcis pityusensis * (Squamata: Lacertidae).
 Molecular Ecology 22, 4829–4841.
 [https://doi.org/10.1111/mec.12443](https://doi.org/10.1111/mec.12443)