DOI: 10.5061/dryad.sn02v6x3q
title: Habitat structure mediates vulnerability to climate change through its effects on thermoregulatory behavior

description: Tropical ectotherms are thought to be especially vulnerable to climate
 change because they are thermal specialists, having evolved in aseasonal
 thermal environments. However, even within the tropics, habitat structure
 can influence opportunities for behavioral thermoregulation. Open (and
 edge) habitats likely promote more effective thermoregulation due to the
 high spatial heterogeneity of the thermal landscape, while forests are
 thermally homogenous and may constrain opportunities for behavioral
 buffering of environmental temperatures. Nevertheless, the ways in which
 behavior and physiology interact at local scales to influence the response
 to climate change are rarely investigated. We examined the thermal ecology
 and physiology of two lizard species that occupy distinct environments in
 the tropics. The brown anole lizard (Anolis sagrei) lives along forest
 edges in The Bahamas, whereas the Panamanian slender anole (Anolis
 apletophallus) lives under the canopy of mature forests in Panama. We
 combined detailed estimates of environmental variation, thermoregulatory
 behavior, and physiology to model the vulnerability of each of these
 species. Our projections suggest that forest-dwelling slender anoles will
 experience severely reduced locomotor performance, activity time, and
 energy budgets as the climate warms over the coming century. Conversely,
 the forest-edge dwelling brown anoles may use behavioral compensation in
 the face of warming, maintaining population viability for many decades.
 Our results indicate that local habitat variation, through its effects on
 behavior and physiology, is a major determinant of vulnerability to
 climate change. When attempting to predict the impacts of climate change
 on a given population, broad-scale characteristics such as latitude may
 have limited predictive power.
description: Please see the methods section of the published manuscript
description: This Neel_Anole_Biotropica_readme.txt file was generated on
 2021-03-29 by Lauren Neel GENERAL INFORMATION
 1. Title of manuscript: Habitat structure mediates vulnerability
 to climate change through its effects on thermoregulatory
 behavior 2. Author Information<br>     A. Lead
 Author Contact Information<br>         Name: Lauren Neel<br>
         Institution: Arizona State University <br>         Address:
 Tempe, AZ, USA<br>         Email: lkneel@asu.edu
     B. Associate or Co-investigator Contact Information<br>
         Name: Mike Logan<br>         Institution: University Nevada
 - Reno<br>         Address: Reno, NV, USA<br>         Email:
 mike.logan1983@gmail.com <br> 3. Date of data
 collection (single date, range, approximate date): Please see Table S1 in
 the manuscript for this detailed information. 4.
 Geographic location of data collection: <br>     1. Great Exuma, The
 Bahamas <br>     2. Gamboa, Panama 5. Information
 about funding sources that supported the collection of the
 data:  Our methods were approved by the Smithsonian
 Tropical Research Institute Institutional Animal Care and Use Committee
 (protocol 2017-0308-2020-A5), Harvard University Institutional Animal Care
 and Use Committee (protocol 26-11), MiAmbiente research permit SE/A-37-19,
 and the BEST commission research permit. Funding for this project was
 provided by a USAID Global Development Research Fellowship awarded to L.
 Neel, a STRI-ASU Collaborative Initiative Research Grant awarded to L.
 Neel and M. Logan, a Smithsonian Institution Biodiversity Genomics
 Postdoctoral Fellowship awarded to M. Logan, a Smithsonian Tropical
 Research Institute Earl S. Tupper Postdoctoral Fellowship awarded to M.
 Logan,  a NERC studentship (NE/L002485/1) awarded to D. Nicholson, a
 Smithsonian Pre-Doctoral Fellowship awarded to D. Nicholson, a STRI
 Short-Term Fellowship awarded to A. Chung, Georgia Southern University
 Graduate Student Organization Professional Development grants awarded to
 A. Chung and J. Curlis, American Museum of Natural History Theodore
 Roosevelt grants awarded to A. Chung and J. Curlis, and a John Templeton
 Foundation grant awarded to J. Losos (the opinions expressed in this
 publication are those of the authors and do not necessarily reflect the
 views of the John Templeton Foundation). The authors thank Lil Camacho,
 Adriana Bilgray, Paola Gomez, and Raineldo Urriola for administrative
 support at STRI.  <br> SHARING/ACCESS
 INFORMATION 1. Licenses/restrictions placed on the
 data: n/a 2. Links to publications that cite or use the
 data: n/a 3. Links to other publicly accessible
 locations of the data: n/a 4. Links/relationships to
 ancillary data sets: n/a 5. Was data derived from
 another source? no <br> DATA & FILE
 OVERVIEW 1. File List:    
  "2017-2018 Panama and Bahamas CTmin UVT.csv" - file containing
 thermal tolerance data<br>   
  "activity_time_model_panama.xlsx" - file containing modeled
 body temperature data for Panama, generated using regression equation from
 present day body temperature (Tb) and operative temperature (Te) data.
 Body temps modeled for a 3C warming event occurring over 100 years.
 Activity is assumed to be possible whenever Tb is below upper voluntary
 temperatures. <br>     "activity_time_model_bahamas.xlsx"
 - file containing modeled body temperature data for The Bahamas, generated
 using regression equation from present day body temperature (Tb) and
 operative temperature (Te) data. Body temps modeled for a 3C warming event
 occurring over 100 years. Activity is assumed to be possible whenever Tb
 is below upper voluntary temperatures. <br>     "Anole Mainland
 Tb.csv" - field active body temperature data<br>   
  "combined mainland data sheet.csv" - file containing Tb and Te
 when captured, thermal tolerance, field data, mass, svl<br>   
  "combined mainland Tpref.csv" - file containing more detailed
 thermal preference data<br>     "metabolism.csv" - file
 containing lizard sex, mass, oxygen consumed, trial temperature, and
 calculated Q10 values<br>     "MR modeling panama.xlsx" -
 file containing modeled body temperature data and oxygen consumption for
 3C warming event, assuming current relationship between metabolism and
 temperature in Panama<br>     "MR modeling bahamas.xlsx" -
 file containing modeled body temperature data and oxygen consumption for
 3C warming event, assuming current relationship between metabolism and
 temperature in The Bahamas<br>     "Sprint projection
 panama.xlsx" - file containing modeled body temperature data and
 sprint speeds for 3C warming event, assuming current relationship between
 locomotor performance and temperature in Panama<br>     "Sprint
 projection bahamas.xlsx" - file containing modeled body temperature
 data and sprint speeds for 3C warming event, assuming current relationship
 between locomotor performance and temperature in The Bahamas<br>   
  "sprint_thermal_sensitivity.csv" - file containing thermal
 sensitivity of locomotor performance data<br>   
  "TbTe_andmass.xlsx" - file containing lizard mass, Tb when
 caught, and average Te when caught. Regression equation from Tb plotted as
 function of Te retained to incorporate thermoregulation when modeling body
 temperatures under 3C warming scenario. <br>   
  "Hourly_Te.xlsx" - file containing summary hourly operative
 environmental data <br>   
  "daily_nightly_summary_Te.xlsx" - daily and nightly summary
 operative environmental data<br>     "monthly weather station
 data bahamas.xlsx" - monthly weather station air temperature data in
 the Bahamas<br>     "monthly weather station data
 panama.xlsx" - monthly weather station air temperature data in
 Panama <br> 2. Relationship between files, if
 important: n/a 3. Additional related data collected
 that was not included in the current data package: no
 4. Are there multiple versions of the dataset? no<br>   
  A. If yes, name of file(s) that was updated: <br>         i. Why
 was the file updated? <br>         ii. When was the file
 updated?  <br> METHODOLOGICAL
 INFORMATION 1. Description of methods used for
 collection/generation of data: <br> Please read the full materials
 and methods section of our manuscript for this detailed
 information.  2. Methods for processing the
 data: <br> Please read the full materials and methods section of our
 manuscript for this detailed information.  3.
 Instrument- or software-specific information needed to interpret the
 data: <br> Model comparisons were conducted with the MUMIN package
 in R version 3.6.2 (R Core Team 2020).  4. Standards
 and calibration information, if appropriate: n/a 5.
 Environmental/experimental conditions: n/a 6. Describe
 any quality-assurance procedures performed on the data: n/a
 7. People involved with sample collection, processing, analysis
 and/or submission: all authors were involved with some combination of
 these items. DATA-SPECIFIC INFORMATION FOR: [2017-2018
 Panama and Bahamas CTmin UVT.csv] 1. Number of
 variables: 4 2. Number of cases/rows: 1909
 3. Variable List: <br> &lt;list variable name(s),
 description(s), unit(s)and value labels as appropriate for
 each&gt;<br> Country - population of individual
 sampled<br> Year - year data collected<br> Ctmin - critical
 thermal minima (C)<br> UVT - upper voluntary temperature
 (C) <br> 4. Missing data codes: <br> N/a -
 means data type were not collected from that individual. 
 5. Specialized formats or other abbreviations used: <br>
 N/a DATA-SPECIFIC INFORMATION FOR:
 [activity_time_model_panama.xlsx]<br> &lt;repeat this section
 for each dataset, folder or file, as appropriate&gt;
 1. Number of variables: 4 2. Number of
 cases/rows: 82 3. Variable List: <br> Hours
 active - total number annual hours where predicted Tb was within preferred
 thermal range (CTmin - UVT) <br> Year - year data modeled<br>
 Daily hours - total number daily hours where predicted Tb was within
 preferred thermal range (CTmin - UVT) <br> % annual hours -
 percentage of annual hours where predicted Tb within preferred thermal
 range <br> 4. Missing data codes: <br> N/a
 - means data type were not collected from that individual. 
 5. Specialized formats or other abbreviations used: <br>
 N/a DATA-SPECIFIC INFORMATION FOR:
 [activity_time_model_bahamas.xlsx]<br> &lt;repeat this section
 for each dataset, folder or file, as appropriate&gt;
 1. Number of variables: 4 2. Number of
 cases/rows: 82 3. Variable List: <br> Hours
 active - total number annual hours where predicted Tb was within preferred
 thermal range (CTmin - UVT) <br> Year - year data modeled<br>
 Daily hours - total number daily hours where predicted Tb was within
 preferred thermal range (CTmin - UVT) <br> % annual hours -
 percentage of annual hours where predicted Tb within preferred thermal
 range <br> 4. Missing data codes: <br> N/a
 - means data type were not collected from that individual. 
 5. Specialized formats or other abbreviations used: <br>
 N/a DATA-SPECIFIC INFORMATION FOR: [Anole Mainland
 Tb.csv]<br> &lt;repeat this section for each dataset, folder or
 file, as appropriate&gt; 1. Number of variables:
 4 2. Number of cases/rows: 1426 3.
 Variable List: <br> Lizard ID - Lizard identifier<br> Date
 capture - date sampled<br> Capture_site - population <br> Tb -
 field active body temperature (C)  4. Missing data
 codes: <br> N/a - means data type were not collected from that
 individual.  5. Specialized formats or other
 abbreviations used: <br> N/a <br>
 DATA-SPECIFIC INFORMATION FOR: [combined mainland data
 sheet.csv]<br> &lt;repeat this section for each dataset, folder
 or file, as appropriate&gt; 1. Number of variables:
 11 2. Number of cases/rows: 2013 3.
 Variable List: <br> Lizard ID - Lizard identifier<br> Country
 - population of individual sampled<br> Year - year data
 collected<br> Timecaptured - time animal captured<br> Tb -
 field active body temp (C)<br> Te - average operative temperature
 (C) at time lizard caught<br> CTmin - lower thermal limit
 (C)<br> UVT - upper voluntary thermal limit (C)<br> Sex - sex
 m/f<br> Mass_g - mass (g)<br> SVL_mm - Snout vent length
 (mm) <br> 4. Missing data codes: <br> N/a -
 means data type were not collected from that individual. 
 5. Specialized formats or other abbreviations used: <br>
 N/a <br> DATA-SPECIFIC INFORMATION FOR: [combined
 mainland Tpref.csv]<br> &lt;repeat this section for each
 dataset, folder or file, as appropriate&gt; 1.
 Number of variables: 17 2. Number of cases/rows:
 120 3. Variable List: <br> Country - population
 of individual sampled<br> Trial date - date of trial<br> Time
 trial began - trial start time<br> Lizard ID - Lizard
 identifier<br> Mass - mass (g)<br> SVL_mm - Snout vent length
 (mm)<br> Sex - sex m/f<br> Tpref_mean - mean thermal
 preference (C)<br> Tpref_median - median thermal preference
 (C)<br> Tpref_min - min thermal preference (C)<br> Tpref_max -
 max thermal preference (C)<br> Tpref_range - range of thermal
 preference (C)<br> Tpref_StdDev - standard deviation of preferred
 temps (C)<br> Tpref_25thq - 25th quartile of thermal preference
 (C)<br> Tpref_75thq - 25th quartile of thermal preference
 (C)<br> IQR - interquartile range (C)<br> TC # - thermocouple
 number <br> 4. Missing data codes: <br> N/a
 - means data type were not collected from that individual. <br> ? -
 means data weren't collected for unknown reasons.
 5. Specialized formats or other abbreviations used: <br>
 N/a <br> DATA-SPECIFIC INFORMATION FOR:
 [metabolism.csv]<br> &lt;repeat this section for each dataset,
 folder or file, as appropriate&gt; 1. Number of
 variables: 9 2. Number of cases/rows: 145
 3. Variable List: <br> Lizard - Lizard identifier<br>
 Country - population of individual sampled<br> Year - year data
 collected<br> Sex - sex m/f<br> Mass - mass (g)<br>
 Total o2 consumed per hourly - total hourly oxygen consumed <br>
 Temp - trial temperature (C)<br> Q10 - calculated Q10
 value<br> LH - low/high temp.. used to code in stats
 <br> 4. Missing data codes: <br> N/a - means data
 type were not collected from that individual.  5.
 Specialized formats or other abbreviations used: <br> N/a
 <br> DATA-SPECIFIC INFORMATION FOR: [MR modeling
 bahamas.xlsx]<br> &lt;repeat this section for each dataset,
 folder or file, as appropriate&gt; 1. Number of
 variables: 7 2. Number of cases/rows: 82
 3. Variable List: <br> Te - average annual operative
 temperature incorporating 3C uniform warming scenario occurring through
 century's end<br> Year - year <br> Predicted Tb -
 predicted body temperature (C) given operative temperature under warming
 scenario, adjusted to incorporate thermoregulation (via function from Tb
 Te regression)<br> Predicted VO2 - predicted oxygen consumption (ml
 O2/ gram animal / time ) using current relationship between metabolism and
 temperature<br> EE - annual energy expenditure
 (kilocalorie/hour)<br> EE - annual energy expenditure
 (kilojoule/hour)<br> EE annual - annual energy expenditure assuming
 animal is active 12 hr/day <br> 4. Missing data
 codes: <br> N/a - means data type were not collected from that
 individual.  5. Specialized formats or other
 abbreviations used: <br> N/a <br>
 DATA-SPECIFIC INFORMATION FOR: [MR modeling panama.xlsx]<br>
 &lt;repeat this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 7
 2. Number of cases/rows: 82 3. Variable
 List: <br> Te - average annual operative temperature incorporating
 3C uniform warming scenario occurring through century's end<br>
 Year - year <br> Predicted Tb - predicted body temperature (C) given
 operative temperature under warming scenario, adjusted to incorporate
 thermoregulation (via function from Tb Te regression)<br> Predicted
 VO2 - predicted oxygen consumption (ml O2/ gram animal / time ) using
 current relationship between metabolism and temperature<br> EE -
 annual energy expenditure (kilocalorie/hour)<br> EE - annual energy
 expenditure (kilojoule/hour)<br> EE annual - annual energy
 expenditure assuming animal is active 12 hr/day
 <br> 4. Missing data codes: <br> N/a - means data
 type were not collected from that individual.  5.
 Specialized formats or other abbreviations used: <br> N/a
 <br> DATA-SPECIFIC INFORMATION FOR: [Sprint projection
 bahamas.xlsx]<br> &lt;repeat this section for each dataset,
 folder or file, as appropriate&gt; 1. Number of
 variables: 3 2. Number of cases/rows: 82
 3. Variable List: <br> Year - year <br> Predicted Tb
 - predicted body temperature (C) given operative temperature under warming
 scenario, adjusted to incorporate thermoregulation (via function from Tb
 Te regression)<br> Predicted relative performance - predicted
 relative locomotor performance (meter/second) using current relationship
 between locomotion and temperature <br> 4.
 Missing data codes: <br> N/a - means data type were not collected
 from that individual.  5. Specialized formats or other
 abbreviations used: <br> N/a DATA-SPECIFIC
 INFORMATION FOR: [Sprint projection panama.xlsx]<br> &lt;repeat
 this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 3
 2. Number of cases/rows: 82 3. Variable
 List: <br> Year - year <br> Predicted Tb - predicted body
 temperature (C) given operative temperature under warming scenario,
 adjusted to incorporate thermoregulation (via function from Tb Te
 regression)<br> Predicted relative performance - predicted relative
 locomotor performance (meter/second) using current relationship between
 locomotion and temperature <br> 4. Missing data
 codes: <br> N/a - means data type were not collected from that
 individual.  5. Specialized formats or other
 abbreviations used: <br> N/a DATA-SPECIFIC
 INFORMATION FOR: [Sprint_thermal_sensitivity.xlsx]<br>
 &lt;repeat this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 6
 2. Number of cases/rows: 65 3. Variable
 List: <br> Lizard # - Lizard identifier<br> Topt - thermal
 optima for locomotor performance (C)<br> Pmax - maximum locomotor
 performance (m/s)<br> Mass - mass (g)<br> Country - population
 of individual sampled 4. Missing data codes: <br>
 N/a - means data type were not collected from that individual. 
 5. Specialized formats or other abbreviations used: <br>
 N/a DATA-SPECIFIC INFORMATION FOR:
 [TbTe_andmass.csv]<br> &lt;repeat this section for each dataset,
 folder or file, as appropriate&gt; 1. Number of
 variables: 8 2. Number of cases/rows: 692
 3. Variable List: <br> Lizard ID - Lizard
 identifier<br> Time - time lizard sampled<br> Date - date
 lizard sampled<br> Mass - mass (g) of lizard<br> Tb - field
 active body temperature (C)<br> Te - operative environmental
 temperature (C) at time lizard caught<br> #OTMS averaged - the
 number of operative temperature models that went in to averaged Te
 value<br> Country - population of individual sampled
 4. Missing data codes: <br> N/a - means data type were not
 collected from that individual.  5. Specialized formats
 or other abbreviations used: <br> N/a
 DATA-SPECIFIC INFORMATION FOR: [Hourly_Te.xlsx]<br>
 &lt;repeat this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 5
 2. Number of cases/rows: 25 3. Variable
 List: <br> Site - location <br> Hour - hour data
 collected<br> Temp median - median operative temperature
 (C)<br> Se - standard error of operative temperature
 4. Missing data codes: <br> N/a - means data type were not
 collected from that individual.  5. Specialized formats
 or other abbreviations used: <br> N/a
 DATA-SPECIFIC INFORMATION FOR:
 [daily_nightly_summary_Te.xlsx]<br> &lt;repeat this section for
 each dataset, folder or file, as appropriate&gt; 1.
 Number of variables: 3 2. Number of cases/rows:
 178 3. Variable List: <br> Te - operative
 temperature (C)<br> Category - min/max/mean<br> Country -
 country sampled <br> 4. Missing data
 codes: <br> N/a - means data type were not collected from that
 individual.  5. Specialized formats or other
 abbreviations used: <br> N/a DATA-SPECIFIC
 INFORMATION FOR: [monthly weather station data bahamas.xlsx]<br>
 &lt;repeat this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 3
 2. Number of cases/rows: 178 3. Variable
 List: <br> Date - date temp taken<br> Temp - weather station
 air temp (C)<br> Month - month data sampled  4.
 Missing data codes: <br> N/a - means data type were not collected
 from that individual.  5. Specialized formats or other
 abbreviations used: <br> N/a DATA-SPECIFIC
 INFORMATION FOR: [monthly weather station data panama.xlsx]<br>
 &lt;repeat this section for each dataset, folder or file, as
 appropriate&gt; 1. Number of variables: 3
 2. Number of cases/rows: 178 3. Variable
 List: <br> Date - date temp taken<br> Temp - weather station
 air temp (C)<br> Month - month data sampled  4.
 Missing data codes: <br> N/a - means data type were not collected
 from that individual.  5. Specialized formats or other
 abbreviations used: <br> N/a <br>