DOI: 10.7291/d1q96h
title: Momma's larvae: Maternal oceanographic experience and larval size influence early survival of rockfishes

description: Identifying factors that affect larval mortality is critical for
 understanding the drivers of fish population dynamics. Although larval
 fish mortality is high, small changes in mortality rates can lead to large
 changes in recruitment. Recent studies suggest maternal provisioning can
 dramatically affect the susceptibility of larvae to starvation and
 predation, the major sources of early-life mortality. We measured otolith
 core width-at-extrusion and validated that this is a proxy for larval
 size-at-extrusion for eight species of rockfishes
 (genus Sebastes) to examine the influence of initial larval size
 on larval growth and survival and to understand how oceanographic
 conditions experienced by gestating females affect larval size (i.e.,
 quality). Otolith core width-at-extrusion was significantly positively
 related to larval rockfish recent growth rate (5/7 species with sufficient
 sample size) and survival (all eight species). This suggests that
 individuals that are larger at extrusion generally grow faster and are
 more likely to survive early life stages. Otolith core width-at-extrusion
 was positively related to higher presence of Pacific Subarctic Upper Water
 and was negatively related to warmer, saline waters at the depths
 gestating mothers inhabited during the months prior to larval collection.
 In addition, otolith core width was larger further from fishing ports,
 possibly because these locations were historically less fished, contained
 more older, larger females, and/or had inherently better habitat quality
 (higher Pacific Subarctic Upper Water) than sites closer to shore. These
 results indicate that the environmental conditions female rockfish
 experience during gestation drive the size of the larvae they produce and
 impact larval growth and survival.
description: <em>Sample collection </em>
 Rockfish larvae were collected during winter CalCOFI cruises from
 1998 to 2013 as described in Thompson et al. (2017). Winter cruises take
 place in January-February with the majority (72–98%) of larval rockfish
 collected in January (Table 1). Bongo nets were deployed at fixed stations
 to 210 m and obliquely towed to the surface at a constant rate (Kramer
 1972; Figure 1).  Plankton from the port-side net was initially preserved
 in 95% ethanol and changed for fresh 95% ethanol after 24 hours. In the
 laboratory, rockfish larvae were sorted and enumerated from stations over
 the continental shelf (inside yellow polygon figure 1). Because most
 rockfish larvae are not morphologically distinguishable to the species
 level, the mitochondrial cytochrome b gene was sequenced from each
 individual larva using DNA extracted from rockfish tissue (see Thompson et
 al. 2017 for details on sequencing), and identities were determined by
 comparing larval sequence to reference sequences from adults (64).  After
 sequencing, larvae were stored individually in vials filled with 100%
 ethanol. <em>Rockfish early life history trait
 measurement</em> We focused our analysis on eight
 rockfish species found throughout the Southern California Bight. Five of
 the species are commercially/recreationally fished
 (<em>S</em>. <em>goodei</em>,
 <em>S</em>. <em>mystinus</em>,
 <em>S</em>. <em>paucispinis</em>,
 <em>S</em>. <em>rufus,</em>
 <em>S</em>. <em>serranoides</em>) and three are
 non-targeted (<em>S</em>. <em>hopkinsi</em>,
 <em>S</em>. <em>jordani</em>,
 <em>S</em>. <em>wilsoni</em>), although these
 non-targets species are occasionally caught as bycatch in other fisheries.
 A total of 5,475 individuals from all 8 target species were collected
 between 1998–2013. Of those, 2021 were selected for otolith extraction (a
 maximum of 20 larvae from each species per year). Larvae were placed in a
 drop of deionized water on a microscope slide under a dissecting
 microscope and basic morphometrics, including total length, were recorded.
 Sagittal otoliths were then extracted from each fish using micro-forceps
 and insect mounting pins. The otoliths were affixed whole to microscope
 slides with a drop of clear fingernail polish. Polishing was not
 necessary, as intact otoliths from larvae (&lt; 20 mm TL) had
 discernable growth increments with no further preparation.
 Otoliths were photographed using a DSLR camera (Nikon D7000)
 mounted to a compound microscope (Leica DMLB) at 1000× magnification.
 Images were analyzed in R using the <em>RFishBC</em> package
 (v0.2.4; Ogle 2022). This package facilitates measuring otolith increment
 widths and estimating ages by applying a known scale to defined
 measurements made on collected images. Otolith images were batched and
 analyzed by year and species.  Otolith core width-at-extrusion (hereafter
 otolith core width) was measured from the center of the core to the
 postrostral edge of the extrusion check (a mark in the otolith that forms
 when the larva is released from the mother). Ages were obtained by
 increment counts beginning at the first growth increment after the
 extrusion check. For quality control, a randomized sub-sample of images
 was examined by an experienced reader twice, with an extended time
 interval between reads. If age estimates differed between the first two
 reads, the otolith photographs were examined for a third time. Samples
 were not used in further analyses if the third reading did not agree with
 one of the first two. Of the 2,021 otoliths aged, 1,502 had agreement
 between reads, and 1,499 were used for this study (3 individuals were
 collected in offshore stations outside of the region of study; Table 1).
 Growth increments were not validated during this study. However, these
 counts were assumed to correspond to daily ages because the formation of
 daily growth increments has been validated for several of these species,
 including <em>S</em>. <em>jordani</em> (66),
 <em>S</em>. <em>paucispinis</em>, and
 <em>S</em>. <em>goodei</em> (67). 
 <em>Oceanographic conditions & Source-water
 analysis </em> We gathered a suite of
 oceanographic data from basin, regional, and station scales to
 characterize the oceanographic conditions pregnant female rockfishes and
 larval rockfishes experienced. Based on previous work (Schroeder et al.
 2018), we hypothesized that the conditions females experience may play an
 important role in determining larval quality. At the basin scale, we
 included the Pacific Decadal Oscillation (PDO), a measure of low-frequency
 sea surface temperature variability throughout the northeast Pacific, the
 Ocean Niño Index (ONI), which measures equatorial upper water temperature
 and is an indicator of El Niño Southern Oscillation State, and the North
 Pacific Gyre Oscillation (NPGO), an index of the strength of equatorward
 flow of the California Current, vertical mixing, and surface salinity,
 nutrient, and chlorophyll concentrations in the southern CCLME (68).
 Regional oceanographic variables included two measures of wind-driven
 upwelling intensity: the Biologically Effective Upwelling Transport Index
 (BEUTI; Jacox et al. 2018) at 33°N, and meridional winds at 33.7°N and
 119.1°W (from NOAA buoy 46025). At the local scale of the CalCOFI stations
 where rockfish were collected, we calculated the distance to the nearest
 fishing port (also a proxy for distance to shore) and two indicators of
 the source waters contributing to water column characteristics: the PSUW
 fraction and Spiciness (Schroeder et al. 2018). Spiciness is a state
 variable and is derived from temperature and salinity, with lower spice
 values characteristic of cooler, fresher waters (Flament 2002). Water-mass
 fraction, at each CalCOFI station and in the upper 500 m, is obtained from
 an optimal multiparameter (OMP) analysis of 3 water masses that comprise
 upper waters in the CCS (Bograd et al. 2019). Spiciness has been used to
 describe changes in interannual variability of rockfish recruitment as
 abundances of pelagic juveniles (the index of recruitment in stock
 assessments; e.g., He and Field 2017) were higher for most species during
 years with low spiciness (Schroeder et al. 2018). An OMP analysis, using
 CalCOFI hydrographic data starting from 1984, quantifies the mixing
 between PSUW, Eastern North Pacific Central Water (ENPCW), and Pacific
 Equatorial Water (PEW) water masses (Bograd et al. 2019). Of the three
 water masses used in the OMP analysis, PSUW is the water mass of subarctic
 origin that is advected into the CCE, with the greatest percentages
 occurring below 100 m. PSUW has the lowest spiciness values, and we used
 the PSUW fraction in our analyses. Spice and PSUW fraction have been
 calculated for all CalCOFI stations used in this study at 5 m depth
 intervals from the surface to 500 m. Profiles of spice and PSUW fraction
 were linearly interpolated on 1 m intervals and values are extracted on
 isopycnals σθ = 25.5, 25.8, 26.1, 26.5, 26.8 kg/m<sup>3</sup>
 covering depths ranging between 50 m to 400 m. We report PSUW fraction as
 a percentage (PSUW%) at a given time (season and year), station, and
 depth.  The combination of all three water masses (PSUW, ENPCW, and PEW)
 will equal to 1. OMP analysis has been calculated for all quarterly
 CalCOFI surveys, however, for this study, only the fall and winter
 quarters are considered to align with rockfish gestation in fall and
 larval rockfish early life in winter. <em>Data
 analysis</em> To determine whether the otolith
 core width provided a proxy for larval fish size-at-extrusion, as has been
 demonstrated in other species (Garrido et al. 2015), we used linear
 regression to examine the amount of variability in length of all
 one-day-old rockfish larvae explained by otolith core width. While we
 didn’t have immediately-extruded (day 0) larvae available, we assumed that
 one-day-old larvae provided an accurate representation of larval
 length-at-extrusion. We examined the relationship
 between timing of extrusion and otolith core width to determine if core
 size was influenced by when larvae were extruded as there is evidence that
 older, larger females reproduce earlier in the reproductive season. We
 again used linear regression with date of extrusion as our explanatory
 variable to test this relationship. We evaluated time
 series of annual, species-specific, average anomalies of larval rockfish
 early life history traits to determine if patterns of otolith core width,
 recent growth, and age-at-capture were consistent across species (Figure
 S1). A time series of life history trait anomalies was calculated by
 taking the spatial average of each trait at all stations for each year and
 species, then subtracting a long-term mean of each trait from the annual
 values. We used Spearman’s rank correlation to evaluate the degree of
 correlation among species for each life history trait (Figures S2–4).
 Additionally, we plotted spatially averaged otolith core width size across
 the entire time series, at each station, and for each species to explore
 the spatial patterns of variability in otolith core width (Fig.
 2). Because we expected interannual and spatial
 variability in maternal and larval oceanographic experience, we calculated
 the mean core width size for each species at each station in each year and
 used partial least squares regression (PLSR; Carrascal et al. 2009)
 analysis with a suite of basin scale, regional, and local oceanographic
 conditions to understand the environmental drivers of otolith core width
 size. All environmental variables, except PSUW% and Spiciness, were
 averaged from September to December to capture the fall conditions adult
 female rockfish experienced while they were gestating. Fall PSUW% and
 spiciness were calculated from fall CalCOFI hydrographic data and were
 calculated along isopycnal layers (25.5, 25.8, 26.1, 26.5, 26.8) that were
 matched to the adult depth range of the different rockfish
 species. PLSR was used to determine how oceanographic
 conditions and maternal investment contribute to larval rockfish recent
 growth rate and survival. Recent growth anomalies were calculated by
 taking the residuals of the relationship between mean growth of the last
 two growth increments and age-at-collection for each larva. Age-at-capture
 was used as a proxy for larval survival because mortality is high during
 this early period; years with older mean ages indicate higher larval
 survival (Garrido et al. 2015). We included the same environmental
 variables as in the core width analysis but averaged these from January to
 February (winter CalCOFI hydrographic surveys for PSUW and Spice) to
 capture the conditions larval rockfish experienced during early life
 before collection in winter CalCOFI cruises. As larval rockfish are
 typically found in the upper 80m (72) we used PSUW% and Spiciness on the
 25.8 isopycnal, which is approximately 60 m deep near shore and 90 m deep
 offshore. We also included mean core width for each species at each
 station and year to examine the influence of larval size at extrusion on
 growth and survival. Similar to the calculation of mean core width, we
 calculated the mean recent growth rate, and mean age at each station in
 each year. In each PLSR analysis we calculated the
 weight squared of each predictor variable to determine the relative
 contribution of each variable to the PLSR components. When the weights of
 each variable are squared, they sum to one, providing the percent
 contribution of each variable to the PLSR component. We used a cutoff of
 PLSR weight<sup>2</sup> &gt; 0.1 to identify variables
 that contributed significantly (more than 10%) to the first PLSR component
 and drivers of variability in maternal investment, larval growth rate, and
 age-at-capture. All otolith analyses were performed in R (v4.1.2; R Core
 Team 2021)) using packages <em>dplyr</em> and
 <em>plsrdepot</em>. Figures were generated using R package
 <em>ggplot2</em>.
description: This README file was generated on 2023-10-20 by Will Fennie GENERAL
 INFORMATION 1. Title of Dataset: Momma's larvae: maternal
 oceanographic experience and larval size influence early survival of
 rockfishes 2. Author Information A. Corresponding Author Contact
 Informatino Name: Will Fennie Insititution: NOAA Alaska Fisheries Science
 Center Address: Seattle, WA Email: will.fennie@noaa.gov 3. Date of
 collection: Fall 1997 - Winter 2013 4. Geographic location of collection:
 Southern California Bight, California, USA 5. Information about funding
 sources that supported the collection of the data: Future Seas (NOAA
 Climate and Fisheries Adaptation Program grant number NA20OAR4310507), and
 CCE LTER (NSF OCE-2224726). SHARING/ACCESS INFORMATION 1.
 Licenses/restrictions placed on the data: CC0 1.0 Universal (CC0 1.0)
 Public Domain 2. Links to publications that cite or use the data:
 Thompson, A. R., Hyde, J. R., Watson, W., Chen, D. C., & Guo, L.
 W. (2016). Rockfish assemblage structure and spawning locations in
 southern California identified through larval sampling. Marine Ecology
 Progress Series, 547, 177-192. Thompson, A. R., Chen, D. C., Guo, L. W.,
 Hyde, J. R., & Watson, W. (2017). Larval abundances of rockfishes
 that were historically targeted by fishing increased over 16 years in
 association with a large marine protected area. Royal Society open
 science, 4(9), 170639. 3. Links to other publicly accessible locations of
 the data: None 4. Links/relationships to ancillary data sets: None 5. Was
 data derived from another source? No A. If yes, list source(s): NA 6.
 Recommended citation for this dataset: Fennie, H. W., Ben‐Aderet, N.,
 Bograd, S. J., Kwan, G. T., Santora, J. A., Schroeder, I. D., &
 Thompson, A. R. (2023). Data fomr: Momma's larvae: Maternal
 oceanographic experience and larval size influence early survival of
 rockfishes. Dryad Digital Repository.https://doi.org/10.7291/D1Q96H. Data
 and file overview: 1. File list: a) age\_size.rda b) calcofi stations with
 date and time 98-13.csv c) Fall\_OC\_growth\_ENV\_PSUW.rda d)
 winter\_OC\_age\_ENV\_PSUW.rda e)
 Mean\_winter\_OC\_dist\_growth\_resid\_goodei\_ENV\_PSUW.rda f)
 Mean\_winter\_OC\_dist\_growth\_resid\_hopkinsi\_ENV\_PSUW.rda g)
 Mean\_winter\_OC\_dist\_growth\_resid\_jordani\_ENV\_PSUW.rda h) No
 dataset for S. mystinus (too few fish &gt;=3 days old - analysis could
 not run with so few data points) i)
 Mean\_winter\_OC\_dist\_growth\_resid\_paucispinis\_ENV\_PSUW.rda j)
 Mean\_winter\_OC\_dist\_growth\_resid\_rufus\_ENV\_PSUW.rda k)
 Mean\_winter\_OC\_dist\_growth\_resid\_serranoides\_ENV\_PSUW.rda l)
 Mean\_winter\_OC\_dist\_growth\_resid\_wilsoni\_ENV\_PSUW.rda m)
 rockfish\_age.rda 2. Relationship between files, if important: None 3.
 Additional related data collected that was not included in the current
 data package: None 4. Are there multiple versions of the dataset? No A. If
 yes, name of file(s) that was updated: NA i. Why was the file updated? NA
 ii. When was the file updated? NA
 #########################################################################
 Data specific information for: age\_size.rda These data are derived from
 NOAA California Cooperative Oceanic Fisheries Investigations cruises in
 the Southern California Bight from 1998-2013. They contain collection
 information, genetic species identification, larval rockfish body and
 otolith measurements. 1. Number of variables: 13 2. Number of cases/rows:
 1502 3. Variable List: <br> \*Year: Year larval rockfish were
 collected (YYYY) <br> \*Month: Month larval rockfish were collected
 (M) <br> \*species: Rockfish species (goodei, hopkinsi, jordani,
 mystinus, paucispinis, rufus, serranoides, wilsoni) <br> \*age:
 larval rockfish age (days) <br> \*rad1: radius to age 1 otolith
 increment (mm) <br> \*rad2: radius to age 2 otolith increment (mm)
 <br> \*TL.mm: Total length (mm) <br> \*Line\_station: Name of
 the station where each fish was collected <br> \*lat: latitude
 (decimal degrees) <br> \*lon: longitude (decimal degrees) <br>
 \*µm\_core: otolith nuclear radius (µm) <br> \*µm\_last2: width of
 last two otolith growth increments (µm) <br> \*inc\_1: first
 increment width (µm) 4. Missing data codes: "NA" (data not
 available) 5. Specialized formats or other abbreviations used: None
 #########################################################################
 Data specific information for: calcofi stations with date and time
 98-13.csv These data contain date and time of the rockfish collections at
 each station. 1. Number of variables: 17 2. Number of cases/rows: 4136 3.
 Variable List: calcofi stations with date and time 98-13.csv <br>
 \*unique.code2: code tying each collection to the cruise name and order of
 occupancy <br> \*unique.code: code tying each station to the line,
 order of occupancy, cruise, and ship <br> \*S\_C: cruise name
 <br> \*S\_SC: ship <br> \*Line: name of the transect (line)
 being sampled <br> \*Station: name of a station on a given line
 being sampled <br> \*longitude: decimal degrees <br>
 \*latitude: decimal degrees <br> \*season: season sampled (winter,
 spring, summer, fall) <br> \*Line\_station: identifier of location
 sampled that includes both the line and station <br> \*S\_OO:
 station order of occupancy on a given cruise <br> \*date: (M/DD/YY)
 <br> \*time: (YYYY-MM-DD time in GMT) <br> \*Month: (M)
 <br> \*day: day in a given month (D) <br> \*Year: (YYYY)
 <br> \*Jday: Day of year 4. Missing data codes: "NA" (data
 not available) 5. Specialized formats or other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR: Fall\_OC\_growth\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for each
 species to examine how oceanographic conditions female rockfish experience
 in fall, during gestion, affect the quality of the larvae they produce in
 winter. These data have mean otolith characteristics averaged for each
 species at a given station in a given year combined with the fall
 oceanographic conditions at basin, regional, and station/depth specific
 level for each species. 1. Number of variables: 23 2. Number of
 cases/rows: 583 3. Variable List: <br> \*Station: name of a station
 on a given line being sampled <br> \*Line: name of the transect
 (line) being sampled <br> \*Year\_larva: year larva was collected
 (YYYY) <br> \*Year\_mom: year that fall oceanographic conditions
 were sampled to capture maternal oceanographic experience (YYYY)
 <br> \*species: Rockfish species (goodei, hopkinsi, jordani,
 mystinus, paucispinis, rufus, serranoides, wilsoni) <br>
 \*mean\_core: mean otolith core width averaged for each species at a given
 station in a given year (µm) <br> \*mean\_distance: distance from
 nearest fishing port (km) <br> \*Year: year (YYYY) that
 oceanographic conditions were sampled (same as Year\_mom) <br>
 \*avgBEUTI: Biologically effective upwelling transport index (BEUTI see
 Jacox et al. 2018. Coastal upwelling revisted: Ekman, Bakun, and improved
 upwelling indices for the U.S. west coast) <br> \*avgONI: Ocean Nino
 Index (ONI a basin scale measurement of sea surface temperature)
 <br> \*avgPDO: Pacific Decadal Oscillation (PDO a measure of
 low-frequency sea surface temperature variability throughout the northeast
 Pacific) \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the
 strength of equatorward flow of the California Current, vertical mixing,
 and surface salinity, nutrient,and chlorophyll concentrations in the
 southern CCLME) <br> \*avgwind: Meridional wind calculated for the
 southern Californa Bight. <br> \*spice\_25.5: Spiciness (is a state
 variable and is derived from temperature and salinity, with lower spice
 values characteristic of cooler, fresher waters) calculated along the 25.5
 potential density isopycnal layer. <br> \*x\_PSUW\_25.5: Percent
 Pacific Subrctic Upper Water at the 25.5 potential density isopycnal
 layer. (PSUW relative contribution of this watermass at a given CTD
 station. PSUW is characterized as low in temperatuer and salinity, high in
 O2, and nutrient rich) <br> \*spice\_25.8: Spiciness calculated
 along the 25.8 potential density isopycnal layer. <br>
 \*x\_PSUW\_25.8: PSUW calculated along the 25.8 potential density
 isopycnal layer. <br> \*spice\_26.1: Spiciness calculated along the
 26.1 potential density isopycnal layer. <br> \*x\_PSUW\_26.1: PSUW
 calculated along the 26.1 potential density isopycnal layer. <br>
 \*spice\_26.5: Spiciness calculated along the 26.5 potential density
 isopycnal layer. <br> \*x\_PSUW\_26.5: PSUW calculated along the
 26.5 potential density isopycnal layer. <br> \*spice\_26.8:
 Spiciness calculated along the 26.8 potential density isopycnal layer.
 <br> \*x\_PSUW\_26.8: PSUW calculated along the 26.1 potential
 density isopycnal layer. 4. Missing data codes: "NA" (data not
 available) 5. Specialized formats or other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR: winter\_OC\_age\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for each
 species to examine how a combination of maternal investment and the
 oceanographic conditions larval rockfish experience in winter after they
 have been extruded affect their survival (using age as a proxy for
 survival). These data have mean otolith characteristics averaged for each
 species at a given station in a given year combined with the winter
 oceanographic conditions at basin, regional, and station/depth specific
 level for each species. 1. Number of variables: 15 2. Number of
 cases/rows: 583 3. Variable List: <br> \*Station: name of a station
 on a given line being sampled <br> \*Line: name of the transect
 (line) being sampled <br> \*Year\_larva: year larva was collected
 (YYYY) <br> \*Year\_mom: year that fall oceanographic conditions
 were sampled to capture maternal oceanographic experience (YYYY)
 <br> \*species: Rockfish species (goodei, hopkinsi, jordani,
 mystinus, paucispinis, rufus, serranoides, wilsoni) <br>
 \*mean\_age: mean age averaged for each species at a given station in a
 given year (days) <br> \*mean\_core: mean otolith core width
 averaged for each species at a given station in a given year (µm)
 <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.5: Spiciness (is a state variable
 and is derived from temperature and salinity, with lower spice values
 characteristic of cooler, fresher waters) calculated along the 25.5
 potential density isopycnal layer. <br> \*x\_PSUW\_25.5: Percent
 Pacific Subrctic Upper Water at the 25.5 potential density isopycnal
 layer. (PSUW relative contribution of this watermass at a given CTD
 station. PSUW is characterized as low in temperatuer and salinity, high in
 O2, and nutrient rich) <br> \*spice\_25.8: Spiciness calculated
 along the 25.8 potential density isopycnal layer. <br>
 \*x\_PSUW\_25.8: PSUW calculated along the 25.8 potential density
 isopycnal layer. 4. Missing data codes: "NA" (data not
 available) 5. Specialized formats or other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_goodei\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for Sebastes
 goodei to examine how a combination of maternal investment and the
 oceanographic conditions larval rockfish experience in winter after they
 have been extruded affect their age corrected recent growth. These data
 have mean otolith characteristics averaged for each species at a given
 station in a given year combined with the winter oceanographic conditions
 at basin, regional, and station/depth specific level for each species. 1.
 Number of variables: 17 2. Number of cases/rows: 28 3. Variable List:
 <br> \*Station: name of a station on a given line being sampled
 <br> \*Line: name of the transect (line) being sampled <br>
 \*Year\_larva: year larva was collected (YYYY) <br> \*Year\_mom:
 year that fall oceanographic conditions were sampled to capture maternal
 oceanographic experience (YYYY) <br> \*species: Rockfish species
 (goodei) <br> \*mean\_resid: mean recent growth-at-age residual for
 S. goodei averaged at each station in a given year <br> \*mean\_age:
 mean age averaged for each species at a given station in a given year
 (days) <br> \*mean\_core: mean otolith core width averaged for each
 species at a given station in a given year (µm) <br>
 \*mean\_distance: distance from nearest fishing port (km) <br>
 \*Year: year (YYYY) that oceanographic conditions were sampled (same as
 Year\_mom) <br> \*avgBEUTI: Biologically effective upwelling
 transport index (BEUTI see Jacox et al. 2018. Coastal upwelling revisted:
 Ekman, Bakun, and improved upwelling indices for the U.S. west coast)
 <br> \*avgONI: Ocean Nino Index (ONI a basin scale measurement of
 sea surface temperature) <br> \*avgPDO: Pacific Decadal Oscillation
 (PDO a measure of low-frequency sea surface temperature variability
 throughout the northeast Pacific) <br> \*avgNPGO: North Pacific Gyre
 Oscillation (NPGO an index of the strength of equatorward flow of the
 California Current, vertical mixing, and surface salinity, nutrient,and
 chlorophyll concentrations in the southern CCLME) <br> \*avgwind:
 Meridional wind calculated for the southern Californa Bight. <br>
 \*spice\_25.8: Spiciness calculated along the 25.8 potential density
 isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW calculated along the
 25.8 potential density isopycnal layer. 4. Missing data codes:
 "NA" (data not available) 5. Specialized formats or other
 abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_hopkinsi\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for Sebastes
 hopkinsi to examine how a combination of maternal investment and the
 oceanographic conditions larval rockfish experience in winter after they
 have been extruded affect their age corrected recent growth. These data
 have mean otolith characteristics averaged for each species at a given
 station in a given year combined with the winter oceanographic conditions
 at basin, regional, and station/depth specific level for each species. 1.
 Number of variables: 17 2. Number of cases/rows: 57 3. Variable List:
 <br> \*Station: name of a station on a given line being sampled
 <br> \*Line: name of the transect (line) being sampled <br>
 \*Year\_larva: year larva was collected (YYYY) <br> \*Year\_mom:
 year that fall oceanographic conditions were sampled to capture maternal
 oceanographic experience (YYYY) <br> \*species: Rockfish species
 (hopkinsi) <br> \*mean\_resid: mean recent growth-at-age residual
 for S. hopkinsi averaged at each station in a given year <br>
 \*mean\_age: mean age averaged for each species at a given station in a
 given year (days) <br> \*mean\_core: mean otolith core width
 averaged for each species at a given station in a given year (µm)
 <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.8: Spiciness calculated along the
 25.8 potential density isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW
 calculated along the 25.8 potential density isopycnal layer. 4. Missing
 data codes: "NA" (data not available) 5. Specialized formats or
 other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_jordani\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for Sebastes
 jordani to examine how a combination of maternal investment and the
 oceanographic conditions larval rockfish experience in winter after they
 have been extruded affect their age corrected recent growth. These data
 have mean otolith characteristics averaged for each species at a given
 station in a given year combined with the winter oceanographic conditions
 at basin, regional, and station/depth specific level for each species. 1.
 Number of variables: 17 2. Number of cases/rows: 58 3. Variable List:
 <br> \*Station: name of a station on a given line being sampled
 <br> \*Line: name of the transect (line) being sampled <br>
 \*Year\_larva: year larva was collected (YYYY) <br> \*Year\_mom:
 year that fall oceanographic conditions were sampled to capture maternal
 oceanographic experience (YYYY) <br> \*species: Rockfish species
 (jordani) <br> \*mean\_resid: mean recent growth-at-age residual for
 S. jordani averaged at each station in a given year <br>
 \*mean\_age: mean age averaged for each species at a given station in a
 given year (days) <br> \*mean\_core: mean otolith core width
 averaged for each species at a given station in a given year (µm)
 <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.8: Spiciness calculated along the
 25.8 potential density isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW
 calculated along the 25.8 potential density isopycnal layer. 4. Missing
 data codes: "NA" (data not available) 5. Specialized formats or
 other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_mystinus\_ENV\_PSUW.rda There were
 insufficient samples to run this analysis for S. mystinus.
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_paucispinis\_ENV\_PSUW.rda These
 data are used to run a partial least squares regression analyses for
 Sebastes paucispinis to examine how a combination of maternal investment
 and the oceanographic conditions larval rockfish experience in winter
 after they have been extruded affect their age corrected recent growth.
 These data have mean otolith characteristics averaged for each species at
 a given station in a given year combined with the winter oceanographic
 conditions at basin, regional, and station/depth specific level for each
 species. 1. Number of variables: 17 2. Number of cases/rows: 29 3.
 Variable List: <br> \*Station: name of a station on a given line
 being sampled <br> \*Line: name of the transect (line) being sampled
 <br> \*Year\_larva: year larva was collected (YYYY) <br>
 \*Year\_mom: year that fall oceanographic conditions were sampled to
 capture maternal oceanographic experience (YYYY) <br> \*species:
 Rockfish species (paucispinis) <br> \*mean\_resid: mean recent
 growth-at-age residual for S. paucispinis averaged at each station in a
 given year <br> \*mean\_age: mean age averaged for each species at a
 given station in a given year (days) <br> \*mean\_core: mean otolith
 core width averaged for each species at a given station in a given year
 (µm) <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.8: Spiciness calculated along the
 25.8 potential density isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW
 calculated along the 25.8 potential density isopycnal layer. 4. Missing
 data codes: "NA" (data not available) 5. Specialized formats or
 other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_rufus\_ENV\_PSUW.rda These data are
 used to run a partial least squares regression analyses for Sebastes rufus
 to examine how a combination of maternal investment and the oceanographic
 conditions larval rockfish experience in winter after they have been
 extruded affect their age corrected recent growth. These data have mean
 otolith characteristics averaged for each species at a given station in a
 given year combined with the winter oceanographic conditions at basin,
 regional, and station/depth specific level for each species. 1. Number of
 variables: 17 2. Number of cases/rows: 31 3. Variable List: <br>
 \*Station: name of a station on a given line being sampled <br>
 \*Line: name of the transect (line) being sampled <br>
 \*Year\_larva: year larva was collected (YYYY) <br> \*Year\_mom:
 year that fall oceanographic conditions were sampled to capture maternal
 oceanographic experience (YYYY) <br> \*species: Rockfish species
 (rufus) <br> \*mean\_resid: mean recent growth-at-age residual for
 S. rufus averaged at each station in a given year <br> \*mean\_age:
 mean age averaged for each species at a given station in a given year
 (days) <br> \*mean\_core: mean otolith core width averaged for each
 species at a given station in a given year (µm) <br>
 \*mean\_distance: distance from nearest fishing port (km) <br>
 \*Year: year (YYYY) that oceanographic conditions were sampled (same as
 Year\_mom) <br> \*avgBEUTI: Biologically effective upwelling
 transport index (BEUTI see Jacox et al. 2018. Coastal upwelling revisted:
 Ekman, Bakun, and improved upwelling indices for the U.S. west coast)
 <br> \*avgONI: Ocean Nino Index (ONI a basin scale measurement of
 sea surface temperature) <br> \*avgPDO: Pacific Decadal Oscillation
 (PDO a measure of low-frequency sea surface temperature variability
 throughout the northeast Pacific) <br> \*avgNPGO: North Pacific Gyre
 Oscillation (NPGO an index of the strength of equatorward flow of the
 California Current, vertical mixing, and surface salinity, nutrient,and
 chlorophyll concentrations in the southern CCLME) <br> \*avgwind:
 Meridional wind calculated for the southern Californa Bight. <br>
 \*spice\_25.8: Spiciness calculated along the 25.8 potential density
 isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW calculated along the
 25.8 potential density isopycnal layer. 4. Missing data codes:
 "NA" (data not available) 5. Specialized formats or other
 abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_serranoides\_ENV\_PSUW.rda These
 data are used to run a partial least squares regression analyses for
 Sebastes serranoides to examine how a combination of maternal investment
 and the oceanographic conditions larval rockfish experience in winter
 after they have been extruded affect their age corrected recent growth.
 These data have mean otolith characteristics averaged for each species at
 a given station in a given year combined with the winter oceanographic
 conditions at basin, regional, and station/depth specific level for each
 species. 1. Number of variables: 17 2. Number of cases/rows: 11 3.
 Variable List: <br> \*Station: name of a station on a given line
 being sampled <br> \*Line: name of the transect (line) being sampled
 <br> \*Year\_larva: year larva was collected (YYYY) <br>
 \*Year\_mom: year that fall oceanographic conditions were sampled to
 capture maternal oceanographic experience (YYYY) <br> \*species:
 Rockfish species (serranoides) <br> \*mean\_resid: mean recent
 growth-at-age residual for S. serranoides averaged at each station in a
 given year <br> \*mean\_age: mean age averaged for each species at a
 given station in a given year (days) <br> \*mean\_core: mean otolith
 core width averaged for each species at a given station in a given year
 (µm) <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.8: Spiciness calculated along the
 25.8 potential density isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW
 calculated along the 25.8 potential density isopycnal layer. 4. Missing
 data codes: "NA" (data not available) 5. Specialized formats or
 other abbreviations used: None
 #########################################################################
 DATA-SPECIFIC INFORMATION FOR:
 Mean\_winter\_OC\_dist\_growth\_resid\_wilsoni\_ENV\_PSUW.rda These data
 are used to run a partial least squares regression analyses for Sebastes
 wilsoni to examine how a combination of maternal investment and the
 oceanographic conditions larval rockfish experience in winter after they
 have been extruded affect their age corrected recent growth. These data
 have mean otolith characteristics averaged for each species at a given
 station in a given year combined with the winter oceanographic conditions
 at basin, regional, and station/depth specific level for each species. 1.
 Number of variables: 17 2. Number of cases/rows: 45 3. Variable List:
 <br> \*Station: name of a station on a given line being sampled
 <br> \*Line: name of the transect (line) being sampled <br>
 \*Year\_larva: year larva was collected (YYYY) <br> \*Year\_mom:
 year that fall oceanographic conditions were sampled to capture maternal
 oceanographic experience (YYYY) <br> \*species: Rockfish species
 (wilsoni) <br> \*mean\_resid: mean recent growth-at-age residual for
 S. wilsoni averaged at each station in a given year <br>
 \*mean\_age: mean age averaged for each species at a given station in a
 given year (days) <br> \*mean\_core: mean otolith core width
 averaged for each species at a given station in a given year (µm)
 <br> \*mean\_distance: distance from nearest fishing port (km)
 <br> \*Year: year (YYYY) that oceanographic conditions were sampled
 (same as Year\_mom) <br> \*avgBEUTI: Biologically effective
 upwelling transport index (BEUTI see Jacox et al. 2018. Coastal upwelling
 revisted: Ekman, Bakun, and improved upwelling indices for the U.S. west
 coast) <br> \*avgONI: Ocean Nino Index (ONI a basin scale
 measurement of sea surface temperature) <br> \*avgPDO: Pacific
 Decadal Oscillation (PDO a measure of low-frequency sea surface
 temperature variability throughout the northeast Pacific) <br>
 \*avgNPGO: North Pacific Gyre Oscillation (NPGO an index of the strength
 of equatorward flow of the California Current, vertical mixing, and
 surface salinity, nutrient,and chlorophyll concentrations in the southern
 CCLME) <br> \*avgwind: Meridional wind calculated for the southern
 Californa Bight. <br> \*spice\_25.8: Spiciness calculated along the
 25.8 potential density isopycnal layer. <br> \*x\_PSUW\_25.8: PSUW
 calculated along the 25.8 potential density isopycnal layer. 4. Missing
 data codes: "NA" (data not available) 5. Specialized formats or
 other abbreviations used: None
 #########################################################################
 Data specific information for: rockfish\_age.rda These data contain date
 and time of the rockfish collections at each station. 1. Number of
 variables: 16 2. Number of cases/rows: 1502 3. Variable List: calcofi
 stations with date and time 98-13.csv <br> \*Line: name of the
 transect (line) being sampled <br> \*Station: name of a station on a
 given line being sampled <br> \*Line\_station: identifier of
 location sampled that includes both the line and station <br>
 \*latitude: decimal degrees <br> \*longitude: decimal degrees
 <br> \*Year: (YYYY) <br> \*Month.x: (M) <br> \*species:
 Rockfish species (goodei, hopkinsi, jordani, mystinus, paucispinis, rufus,
 serranoides, wilsoni) <br> \*age: larval rockfish age (days)
 <br> \*µm\_core: otolith nuclear radius (µm) <br> \*µm\_last2:
 width of last two otolith growth increments (µm) <br> \*date:
 (M/DD/YYYY) <br> \*time: (YYYY-MM-DD time in GMT) <br> \*day:
 day in a given month (d) <br> \*Jday: Day of year (D) <br>
 \*DOB: Date of birth (D) 4. Missing data codes: "NA" (data not
 available) 5. Specialized formats or other abbreviations used: None