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Navarro, MO, Parnell PE, Levin LA.  2018.  Essential market squid (Doryteuthis opalescens) embryo habitat: A baseline for anticipated ocean climate change. Journal of Shellfish Research. 37:601-614.   10.2983/035.037.0313   AbstractWebsite

The market squid Doryteuthis opalescens deposits embryo capsules onto the continental shelf from Baja California to southern Alaska, yet little is known about the environment of embryo habitat. This study provides a baseline of environmental data and insights on factors underlying site selection for embryo deposition off southern California, and defines current essential embryo habitat using (1) remotely operated vehicle-supported surveys of benthos and environmental variables, (2) SCUBA surveys, and (3) bottom measurements of T, S, pH, and O-2. Here, embryo habitat is defined using embryo capsule density, capsule bed area, consistent bed footprint, and association with [O-2] and pH (pCO(2)) on the shelf. Spatial variation in embryo capsule density and location appears dependent on environmental conditions, whereas the temporal pattern of year-round spawning is not. Embryos require [O-2] greater than 160 mu mol and pH(T) greater than 7.8. Temperature does not appear to be limiting (range: 9.9 degrees C-15.5 degrees C). Dense embryo beds were observed infrequently, whereas low-density cryptic aggregations were common. Observations of dense embryo aggregation in response to shoaling of low [O-2] and pH indicate habitat compression. Essential embryo habitat likely expands and contracts in space and time directly with regional occurrence of appropriate O-2 and pH exposure. Embryo habitat will likely be at future risk of compression given secular trends of deoxygenation and acidification within the Southern California Bight. Increasingly localized and dense spawning may become more common, resulting in potentially important changes in market squid ecology and management.

Navarro, MO, Bockmon EE, Frieder CA, Gonzalez JP, Levin LA.  2014.  Environmental pH, O-2 and capsular effects on the geochemical composition of statoliths of embryonic squid Doryteuthis opalescens. Water. 6:2233-2254.   10.3390/w6082233   AbstractWebsite

Spawning market squid lay embryo capsules on the seafloor of the continental shelf of the California Current System (CCS), where ocean acidification, deoxygenation and intensified upwelling lower the pH and [O-2]. Squid statolith geochemistry has been shown to reflect the squid's environment (e. g., seawater temperature and elemental concentration). We used real-world environmental levels of pH and [O-2] observed on squid-embryo beds to test in the laboratory whether or not squid statolith geochemistry reflects environmental pH and [O-2]. We asked whether pH and [O-2] levels might affect the incorporation of element ratios (B:Ca, Mg:Ca, Sr:Ca, Ba:Ca, Pb:Ca, U:Ca) into squid embryonic statoliths as (1) individual elements and/or (2) multivariate elemental signatures, and consider future applications as proxies for pH and [O-2] exposure. Embryo exposure to high and low pH and [O-2] alone and together during development over four weeks only moderately affected elemental concentrations of the statoliths, and uranium was an important element driving these differences. Uranium: Ca was eight-times higher in statoliths exposed to low pHT (7.57-7.58) and low [O-2] (79-82 mu than those exposed to higher ambient pHT (7.92-7.94) and [O-2] (241-243 mu In a separate experiment, exposure to low pHT (7.55-7.56) or low [O-2] (83-86 mu yielded elevated U:Ca and Sr:Ca in the low [O-2] treatment only. We found capsular effects on multiple elements in statoliths of all treatments. The multivariate elemental signatures of embryonic statoliths were distinct among capsules, but did not reflect environmental factors (pH and/or [O-2]). We show that statoliths of squid embryos developing inside capsules have the potential to reflect environmental pH and [O-2], but that these "signals" are generated in concert with the physiological effects of the capsules and embryos themselves.