Factorial field manipulation reveals multi-stressor effects on a critical habitat-forming shellfish
Data files
Oct 03, 2025 version files 131.88 KB
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DayCarbChem_Mussel.csv
11.03 KB
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DayCarbChemProjected.csv
11.87 KB
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DetidedTemp.csv
16.04 KB
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MS_AvgShellCorr_Inner_update.csv
729 B
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MS_AvgShellCorr_Outer_update.csv
671 B
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Mussel_Biomass_Jeb.csv
2.39 KB
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NightCarbChem_Mussel.csv
17.50 KB
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NightCarbChemProjected.csv
11.94 KB
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pH_Meta.csv
9.93 KB
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README.md
19.51 KB
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Shell_Corrosion_Inner.csv
3.51 KB
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Shell_Corrosion_Outer.csv
3.52 KB
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ShellStrength_CL.csv
3.29 KB
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Sitka_Mussel_CLShells.csv
4.76 KB
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TidePool_Propertities_Mussels.csv
664 B
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WaterMeta_DIC_Musselsl.csv
14.51 KB
Abstract
Ocean acidification and warming are expected to have negative impacts on marine organisms, and particularly on shell-building species. Acidification and warming may operate independently or interactively, amplifying or mitigating impacts on individuals. Previous results have primarily come from lab studies of single species, yet these climate stressors are occurring within naturally dynamic systems with high abiotic and biotic variability. As a result, the impacts of these stressors in situ remains poorly understood. We conducted a 6-month field manipulation to determine the effects of ocean acidification and warming on a habitat-forming shellfish, the Pacific blue mussel (Mytilus trossulus), in a dynamic coastal system. Twenty tide pools were factorially manipulated, including unmanipulated control, CO2 added, warmed, and combined CO2 added and warmed treatments. We measured mussel shell thickness, strength, and corrosion at 0, 3, and 6 months of exposure to treatment conditions. CO2 addition led to a decrease in shell thickness and strength and an increase in shell corrosion. However, we also detected an increase in shell strength for mussels exposed to both CO2 addition and warming. These findings indicate that ocean acidification negatively impacted shellfish in situ and that these effects might be mitigated when exposed concurrently to moderate warming, leading to an interactive effect of acidification and warming on this critical habitat-forming shellfish.
This file contains the information about the datasets used in Rangel et al. 2025 where we conducted a 6-month long field manipulation of pH and temperature in 20 tide pools in Sitka, Alaska and measured effects on the mussel Mytilus trossulus shell traits and function. We measured mussel shell thickness, strength, and corrosion.
File names: DayCarbChem_Mussel.csv
- Sample ID: Number of sample in collection sequence
- Date: Date sample was collected
- Month: Month sample was collected
- Year: Year sample was collected
- DayNight: Whether the sample was collected during daytime or nighttime
- OnOff: On means climate manipulations were on and off means when climate manipulations were turned off
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: Indicates whether sample underwent a warmed treatment
- CO2YN: Indicates whether sample underwent a CO2 added treatment (Y = Yes ; N = No)
- Time.water.collected: The actual time sample was collected from tide pools
- Temp: Handheld temperature data (°C)
- Hanna pH: uncorrected handled tide pool pH (mV)
- Salinity: salinity of tide pool (ppt)
- DO: Dissolved oxygen of tide pool (mg/L)
- Light: Value of light from hand held instrument (umol/m/s)
- HannapH_tot: pH of tide pool corrected (total pH units)
- SpecpH_tot: Raw Spec value for pH (total units)
- Raw_TA: Uncorrected total alkalinity values (mmol kg-1)
- Corrected_TA: Salinity corrected Total alkalinity (mmol kg-1)
- Calcite: Saturation state of calcite from CO2sys
- Aragonite: Saturation state of aragonite from CO2sys
- Volume: Volume of water in the pool (L)
- seacarbCO2: CO2 concentration calculated by seacarb packing in R (mmol kg-1)
- fCO2: "potential" fCO2, CO2 fugacity computed at potential temperature and atmospheric pressure (μatm)
- pCO2:"in situ" pCO2, CO2 partial pressure computed at in situ temperature and total pressure (atm + hydrostatic) (μatm)
- fCO2pot: "in situ" fCO2, CO2 fugacity computed at in situ temperature and total pressure (atm + hydrostatic) (μatm)
- HCO3: HCO3 concentration (mmol kg-1)
- CO3: CO3 concentration (mmol kg-1)
- DIC: Dissolved inorganic carbon concentration (mmol kg-1)
- ALK: total alkalinity from seacarb (mmol kg-1)
- OmegaAragonite: seacarb saturation state of aragonite
- OmegaCalcite: seacarb saturation state of calcite
File Name: NightCarbChem_Mussel.csv
- Sample: Number of sample in collection sequence (ID)
- Date: Date sample was collected
- Month: Month sample was collected
- Year: Year sample was collected
- DayNight: Whether the sample was collected during daytime or nighttime
- OnOff: On means climate manipulations were on and off means when climate manipulations were turned off
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Completed: If there is a T1, T2, T3 water sample Y = Yes, N = No
- MissingT: Indicated is there was a missing Timepoint (T1, T2, T3)
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: Indicates whether sample underwent a warmed treatment
- CO2YN: Indicates whether sample underwent a CO2 added treatment (Y = Yes ; N = No)
- SA: Surface area calculated for the pools (m2)
- Volume: Volume of water in tide pools (L)
- DeltaVolume: Change in pool volume with each water collection timepoint through low tide (L)
- Time.water.collected: The actual time sample was collected from tide pools
- Time.water.collected2: The actual time sample was collected from tide pools
- Temp: Handheld temperature data (°C)
- Hanna pH raw: uncorrected handled tide pool pH (mV)
- Calcite: Saturation state of calcite from CO2sys
- Aragonite: Saturation state of aragonite from CO2sys
- Salinity: salinity of tide pool (ppt)
- DO: Dissolved oxygen of tide pool (mg/L)
- Light: Value of light from hand held instrument (umol/m/s)
- Raw_TA: Uncorrected total alkalinity values (mmol kg-1)
- Corrected_TA: Salinity corrected Total alkalinity (mmol kg-1)
- HannapH: pH of tide pool corrected (total pH units)
- SpecpH_tot: Raw Spec value for pH (total units)
- Raw.TA: Uncorrected total alkalinity values (mmol kg-1)
- CorrectedTA: Salinity corrected Total alkalinity (mmol kg-1)
- CorrectedNN: Corrected nutrients (umol/L)
- CorrectedNitrite: Corrected nitrite (umol/L)
- CalculatedNitrate: Calculated nitrate (umol/L)
- Phosphate: Phosphate value (umol/L)
- NH4: Ammonium (umol/L)
- Date_Time: Date and time sample was collected
- TADeltaTime: Change in TA over the sampling period (mmol kg-1)
- DeltaTime: Change in time over the sampling period
- DeltaNN: Change in corrected nutrients over time (umol/L)
- Phosphate: Change in phosphate value over time (umol/L)
- DeltaNH4: Change in ammonium value over time (umol/L)
- DeltaTA_N_Norm: Change in total alkalinity nutrient corrected over time that is normalized (mmol kg-1)
- NEC.mmol.m2.hr: Net ecosystem calcification (mmol m2 hr)
- seacarbCO2: CO2 concentration calculated by seacarb packing in R (mmol kg-1)
- fCO2: "potential" fCO2, CO2 fugacity computed at potential temperature and atmospheric pressure (μatm)
- pCO2: "in situ" pCO2, CO2 partial pressure computed at in situ temperature and total pressure (atm + hydrostatic) (μatm)
- fCO2pot: "in situ" fCO2, CO2 fugacity computed at in situ temperature and total pressure (atm + hydrostatic) (μatm)
- HCO3: HCO3 concentration (mmol kg-1)
- CO3: CO3 concentration (mmol kg-1)
- DIC: Dissolved inorganic carbon concentration (mmol kg-1)
- ALK: total alkalinity from seacarb (mmol kg-1)
- OmegaAragonite: seacarb saturation state of aragonite
- OmegaCalcite: seacarb saturation state of calcite
File names: DayCarbChemProjected.csv and NightCarbChemProjected.csv
- SampleID: Number of sample in collection sequence (ID)
- Date: Date sample was collected
- Month: Month sample was collected
- Year: Year sample was collected
- DayNight: Whether the sample was collected during daytime or nighttime
- OnOff: On means climate manipulations were on and off means when climate manipulations were turned off
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean]
- Completed: If there is a T1, T2, T3 water sample Y = Yes, N = No
- MissingT: Indicated is there was a missing Timepoint (T1, T2, T3)
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: Indicates whether sample underwent a warmed treatment
- CO2YN: Indicates whether sample underwent a CO2 added treatment (Y = Yes ; N = No)
- SA: Surface area calculated for the pools (m2)
- Volume: Volume of water in tide pools (L)
- DeltaVolume: Change in pool volume with each water collection timepoint through low tide (L)
- Time.water.collected: The actual time sample was collected from tide pools
- Time.water.collected2: The actual time sample was collected from tide pools
- Temp: Handheld temperature data (°C)
- Hanna pH raw: uncorrected handled tide pool pH (mV)
- Salinity: salinity of tide pool (ppt)
- DO: Dissolved oxygen of tide pool (mg/L)
- Light: Value of light from hand held instrument (umol/m/s)
- HannapH: pH of tide pool corrected (total pH units)
- Raw.TA: Uncorrected total alkalinity values (mmol kg-1)
- CorrectedTA: Salinity corrected Total alkalinity (mmol kg-1)
- CorrectedNN: Corrected nutrients (umol/L)
- CorrectedNitrite: Corrected nitrite (umol/L)
- CalculatedNitrate: Calculated nitrate (umol/L)
- Phosphate: Phosphate value (umol/L)
- NH4: Ammonium (umol/L)
- pH: pH calculated from seacarb (total units)
- seacarbCO2: CO2 concentration calculated by seacarb packing in R (mmol kg-1)
- fCO2: "potential" fCO2, CO2 fugacity computed at potential temperature and atmospheric pressure (μatm)
- HCO3: HCO3 concentration (mmol kg-1)
- CO3: CO3 concentration (mmol kg-1)
- DIC: Dissolved inorganic carbon concentration (mmol kg-1)
- ALK: total alkalinity from seacarb (mmol kg-1)
- OmegaAragonite: seacarb saturation state of aragonite
- OmegaCalcite: seacarb saturation state of calcite
File Name: DetidedTemp.csv
- Month: Month sample was collected
- Timepoint: Timepoint of exposure to climate manipulations when mussels were collected (0, 3, 6)
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- DayNight: Whether the sample was collected during daytime or nighttime
- Minimum: Minimum of pool temperatures (°C)
- Maximum: Maximum of pool temperatures (°C)
- Range: Range of pool temperatures (°C)
- Median: Median of pool temperatures (°C)
- Average: Average of pool temperatures (°C)
- DailyAverage: Daily average of pool temperatures (°C)
- Max90Percentile: 90th percentile maximum of pool temperatures (°C)
- Max95Percentile: 95th percentile maximum of pool temperatures (°C)
- Max99Percentile: 99th percentile maximum of pool temperatures (°C)
- Min10Percentile: 10th percentile minimum of pool temperatures (°C)
- DailyMaximum: Daily maximum of pool temperatures (°C)
- Daily90MaxPercentile: Daily 90th percentile maximum of pool temperatures (°C)
- Daily95MaxPercentile: Daily 95th percentile maximum of pool temperatures (°C)
- Daily99MaxPercentile: Daily 99th percentile maximum of pool temperatures (°C)
- DailyMaximumPercentile: Daily Maximum of pool temperatures (°C)
- DailyMinimum: Daily minimum of pool temperatures (°C)
- DailyRange: Daily range of pool temperatures (°C)
- Volume: Volume of water in pools (L)
File Name: MS_AvgShellCorr_Inner_update.csv and MS_AvgShellCorr_Outer_update.csv
- Month: Month sample was collected
- Timepoint: Timepoint of exposure to climate manipulations when mussels were collected (0, 3, 6)
- Treatment: Climate manipulation treatment name
- mean: Mean of shell corrosion for inner or outer for each treatment
- n: Number of shells used in the calculation
- sd: Standard deviation of mean inner and outer shell corrosion
- se: Standard error of mean inner and outer shell corrosion
File Name: Mussel_Biomass_Jeb.csv
- Month: Month sample was collected
- Timepoint: Timepoint of exposure to climate manipulations when mussels were collected (0, 3, 6)
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Pool Volume: Volume of water in pools (L)
- Percent Cover: Percent cover of mussels in the pools (%)
- Surface Area: Surface area of the pools (m2)
- WetBiomass: g L-1
- Norm Biomass: g L-1
File Name: pH_Meta.csv
- Sample ID: Number of sample in collection sequence
- Date: Date sample was collected
- Month: Month sample was collected
- Year: Year sample was collected
- DayNight: Whether the sample was collected during daytime or nighttime
- OnOff: On means climate manipulations were on and off means when climate manipulations were turned off
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- Time.water.collected: The actual time sample was collected from tide pools
- Temp: Handheld temperature data (°C)
- Hanna pH: uncorrected hanna tide pool pH (mV)
- Salinity: salinity of tide pool (ppt)
- DO: Dissolved oxygen of tide pool (mg/L)
- Light: Value of light from hand held instrument (umol/m/s)
- HannapH_tot: pH of tide pool corrected (total pH units)
- Raw_TA: Uncorrected total alkalinity values (mmol kg-1)
- Corrected_TA: Salinity corrected Total alkalinity (mmol kg-1)
- Volume: Volume of water in tide pools (L)
File Name: Sitka_Mussel_CLShells.csv
- Date_Measured: Date shell trait measured
- Month: Month sample was collected
- Timepoint: Timepoint of exposure to climate manipulations when mussels were collected (0, 3, 6)
- Species: Type of species that was collected (mussel)
- Mussel_ID: Identification of mussel number
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- Total_L: total length of mussel shell from lip to base (mm)
- Side: Either left or right valve that was measured (1 = right, 2=left)
- Final_LipA: Averaged shell thickness measurements at the lip of shell (mm)
- Final_MidB: Averaged shell thickness measurements at middle point of shell (mm)
- Final_BaseC: Averaged shell thickness measurements at base of shell (mm)
- StandardA: Total length standardized lip shell thickness (mm)
- StandardB: Total length standardized middle shell thickness (mm)
- Standard C: Total length standardized base shell thickness (mm)
- ThreeTrials: Y is mussels were collected from these pool at all three timepoints - N if mussels were not collected at all 3 timepoints
- Volume: Volume of water in tide pools (L)
File Name: ShellStrength_CL.csv
- ID: Mussel Identification
- Pool: Tide pool number where sample was collected
- Month: Month sample was collected
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- Shell_ID: Shell identification
- Depth: Depth of mussel shell measured (mm)
- Length: Total length of mussel shell (lip to base) measured (mm)
- Width: Width of mussel shell measured (mm)
- Mass: Weight of dried mussel shell (g)
- Strength: Raw strength value measured from Instron force test (N)
- Strength.Stand: Total length standardized strength value (N)
- Valve: Right (R) or left (L) value was used in strength test
- ThreeTrials: Whether or not this pool was collected at 3 timepoints (Y = yes, N = No)
- Volume: Volume of water in tide pools (L)
File Name: Shell_Corrosion_Inner.csv
- Month: Month sample was collected
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Mussel.ID: Mussel Identification
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- Valve: Either left (L) or right (R) valve used in measurement
- Side: Either inner (In) or outer (Out) shell measured
- ThreeTrials: Whether or not this pool was collected at 3 timepoints (Y = yes, N = No)
- Outer_TA: Total outer shell area measured (mm)
- Corroded_Area: Total corroded shell area (mm)
- Uncorroded_Area: Total uncorroded shell area (mm)
- Per_Corroded: Percent of corroded shell area calculated (%)
- Per_Uncorroded: Percent of uncorroded shell area (%)
- SL: Total length of mussel shell valve (mm)
- Gam_Corr: Gamma corrected mussel percent corroded (%)
- Volume: Volume of water in tide pools (L)
File Name: Shell_Corrosion_Outer.csv
- Month: Month sample was collected
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean.
- Mussel.ID: Mussel Identification
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: If the pool had a warming treatment (H = Heat, N = No Heat)
- CO2: If the pool had a CO2 treatment (C = CO2, N = No CO2)
- Valve: Either left (L) or right (R) valve used in measurement
- Side: Either inner (In) or outer (Out) shell measured
- ThreeTrials: Whether or not this pool was collected at 3 timepoints (Y = yes, N = No)
- Outer_TA: Total outer shell area measured (mm)
- Corroded_Area: Total corroded shell area (mm)
- Uncorroded_Area: Total uncorroded shell area (mm)
- Per_Corroded: Percent of corroded shell area calculated (%)
- Per_Uncorroded: Percent of uncorroded shell area (%)
- SL: Total length of mussel shell valve (mm)
- Volume: Volume of water in tide pools (L)
File Name: TidePool_Propertities_Mussels.csv
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Max Depth: Depth of the pool (cm)
- Perimeter: Perimeter of the pool (cm)
- Volume: Volume of water in tide pools (L)
- Interior surface area: Number of squares that were counted to calculate inner pool surface area (squares)
- Tide Height: Location of the pool on the shoreline from waterline to high tide (m)
- A: Surface area calculated for the pools (m2)
File Name: WaterMeta_DIC_Musselsl.csv
- Sample: Number of sample in collection sequence (ID)
- Date: Date sample was collected
- Month: Month sample was collected
- Year: Year sample was collected
- DayNight: Whether the sample was collected during daytime or nighttime
- OnOff: On means climate manipulations were on and off means when climate manipulations were turned off
- Timepoint: Time point during each tide when sample was taken (1-3) with 3 being the time point after the longest time of separation from the ocean
- Completed: If there is a T1, T2, T3 water sample Y = Yes, N = No
- MissingT: Indicated is there was a missing Timepoint (T1, T2, T3)
- Pool: Tide pool number where sample was collected
- Treatment: Climate manipulation treatment name
- Heat: Indicates whether sample underwent a warmed treatment
- CO2YN: Indicates whether sample underwent a CO2 added treatment (Y = Yes ; N = No)
- SA: Surface area calculated for the pools (m2)
- Volume: Volume of water in tide pools (L)
- DeltaVolume: Change in pool volume with each water collection timepoint through low tide (L)
- Time.water.collected: The actual time sample was collected from tide pools
- Temp: Handheld temperature data (°C)
- Hanna pH raw: uncorrected handled tide pool pH (mV)
- Salinity: salinity of tide pool (ppt)
- DO: Dissolved oxygen of tide pool (mg/L)
- Light: Value of light from hand held instrument (umol/m/s)
- HannapH: pH of tide pool corrected (total pH units)
- Raw.TA: Uncorrected total alkalinity values (mmol kg-1)
- CorrectedTA: Salinity corrected Total alkalinity (mmol kg-1)
- CorrectedNN: Corrected nutrients (umol/L)
- CorrectedNitrite: Corrected nitrite (umol/L)
- CalculatedNitrate: Calculated nitrate (umol/L)
- Phosphate: Phosphate value (umol/L)
- NH4: Ammonium (umol/L)
Datasets Include:
- DayCarbChem.csv
- DayCarbChemProjected.csv
- DetidedTemp.csv
- MS_AvgShellCorr_Inner_update.csv
- MS_AvgShellCorr_Outer_update.csv
- Mussel_Biomass_Jeb.csv
- NightCarbChem_Mussel.csv
- NightCarbChemProjected.csv
- pH_Meta.csv
- Shell_Corrosion_Inner.csv
- Shell_Corrosion_Outer.csv
- ShellStrength_CL.csv
- Sitka_Mussel_CLShells.csv
- TidePool_Propertities_Mussels.csv
- WaterMeta_DIC_Musselsl.csv
Temperature Data:
Temperature data was collected at the tide pool level using Onset ® HOBO TidbiT temperature loggers (±0.2 accuracy) that recorded temperature consecutively every 5 min from March to September 2019. Temperature data were summarized for the 28-day periods preceding each collection period (March, July, and September). For each tidepool, we calculated the following thermal parameters: minimum, mean daily minimum, 10th percentile, median, average, range, mean daily range, mean daily average, maximum, mean daily maximum, 90th percentile, mean daily 90th percentile, 95th percentile, 99th percentile, mean daily 90th percentile temperatures, mean daily 95th percentile temperatures, mean daily 99th percentile temperatures. All temperature data was separated into low day and nighttime tides based on known shore heights and observed tide data from Sitka Harbor, Baranof Island, Sitka, Alaska (tbone tides; https://tide.arthroinfo.org/).
Environmental Tide Pool Data:
Tide pool parameters were collected using handheld instruments and total alkalinity was determined at UCSC.
Shell Trait Data:
All shell trait data were measured within the laboratory.
Carbonate Chemistry Variables:
pCO2, DIC, and saturation states of calcite and aragonite were calculated using the R package seacarb.
- Rangel, Racine; Bracken, Matthew; Kroeker, Kristy et al. (2025). Factorial field manipulation reveals multi-stressor effects on a critical habitat-forming shellfish. Zenodo. https://doi.org/10.5281/zenodo.8125694
- Rangel, Racine; Bracken, Matthew; Kroeker, Kristy et al. (2025). Factorial field manipulation reveals multi-stressor effects on a critical habitat-forming shellfish. Zenodo. https://doi.org/10.5281/zenodo.8125693
- Rangel, Racine E.; Bracken, Matthew E. S.; Kroeker, Kristy J. et al. (2025). Factorial field manipulation reveals CO2 and temperature effects on a critical habitat-forming shellfish. Journal of Experimental Biology. https://doi.org/10.1242/jeb.250787
