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Fodrie, FJ, Becker BJ, Levin LA, Gruenthal K, McMillan PA.  2011.  Connectivity clues from short-term variability in settlement and geochemical tags of mytilid mussels. Journal of Sea Research. 65:141-150.   10.1016/j.seares.2010.09.001   AbstractWebsite

The use of geochemical tags in calcified structures of fish and invertebrates is an exciting tool for investigating larval population connectivity. Tag evaluation over relatively short intervals (weeks) may detect environmental and ecological variability at a temporal scale highly relevant to larval transport and settlement. We collected newly settled mussels (Mytilus californianus and M. galloprovincialis) weekly during winter/spring of 2002 along the coast of San Diego, CA, USA, at sites on the exposed coast (SIO) and in a protected coastal bay (HI), to investigate temporal patterns of geochemical tags in mussel shells. Analyses of post-settlement shell via LA-ICP-MS revealed statistically significant temporal variability for all elements we examined (Mg, Mn, Cu, Sr, Cd, Ba, Pb and U). Despite this, our ability to distinguish multielemental signatures between sites was largely conserved. Throughout our 13-week study, SIO and HI mussels could be chemically distinguished from one another in 78-87% of all cases. Settlement varied between 2 and 27 settlers grambyssus(-1) week(-1) at 510 and HI, and both sites were characterized by 2-3 weeks with "high" settlement. Geochemical tags recorded in early larval shell of newly settled mussels differed between "high" and "low" settlement weeks at both sites (MANOVA), driven by Mg and Sr at SIO (p = 0.013) and Sr, Cd, Ba and Pb at HI (p < 0.001). These data imply that shifts in larval sources or transport corridors were responsible for observed settlement variation, rather than increased larval production. In particular, increased settlement at HI was observed concurrent with the appearance of geochemical tags (e.g., elevated Cd), suggesting that those larvae were retained in upwelled water near the mouth of the bay. Such shifts may reflect short-term changes in connectivity among sites due to altered transport corridors, and influence the demography of local populations. (C) 2010 Elsevier B.V. All rights reserved.

Fodrie, FJ, Levin LA.  2008.  Linking juvenile habitat utilization to population dynamics of California halibut. Limnology and Oceanography. 53:799-812.   10.4319/lo.2008.53.2.0799   AbstractWebsite

We investigated the nursery role of four coastal ecosystems for the California halibut (Paralichthys californicus) using the following metrics: (1) contribution in producing the fish that advance to older age classes, (2) connectivity of coastal systems resulting from migration of fish from juvenile to subadult habitats, and (3) effect of nursery habitat usage and availability on subadult population size, specifically evaluating the concentration hypothesis. Potential nurseries were grouped using a robust classification scheme that segregated exposed, bay, lagoon, and estuarine environments. Assignment of nursery origins for individual subadult fish via elemental fingerprinting indicated that exposed coasts, bays, lagoons, and estuaries contributed 31%, 65%, 1%, and 3% of advancing juvenile halibut during 2003, versus 49%, 33%, 16%, and 2% during 2004, respectively. These results were remarkably similar to "expected'' nursery contributions derived from field surveys, suggesting that in this system juvenile distributions were a good indicator of unit-area productivity of juvenile habitats and that density-dependent mechanisms during the juvenile phase did not regulate recruitment pulses. Elemental fingerprinting also demonstrated that individuals egressing from bays did not migrate far from their nursery origins (, 10 km), resulting in reduced connectivity along the 110-km study region over the timescale of approximately one generation. Consequently, we observed considerably higher subadult densities at sites near large bays, while populations distant from large bays appeared to be more influenced by nursery habitat limitation. Over large (similar to 100 km) scales, the location and availability of nursery habitat alternatives had significant effects on the population dynamics of an important member of the ichthyofaunal community of southern California.

Fodrie, FJ, Levin LA, Lucas AJ.  2009.  Use of population fitness to evaluate the nursery function of juvenile habitats. Marine Ecology-Progress Series. 385:39-49.   10.3354/meps08069   AbstractWebsite

Juveniles of many fish and invertebrate species are able to select among a diverse portfolio of nursery habitat alternatives. Environmental heterogeneity among these habitats generates variation in the vital rates of young individuals that may influence overall population dynamics. Therefore, understanding how these habitat options affect population fitness is crucial for identifying habitats that widen bottlenecks in early life histories and promote population persistence. We used cohort analyses and demographic models to explore the population-level consequences of habitat selection by juvenile California halibut Paralichthys californicus in southern California, focusing on population growth rate (lambda) as a measure of fitness. Although alternative juvenile habitats (exposed coast and coastal embayments) could contribute an approximately equal number of recruits to the adult stock, positive overall population growth (lambda > 1) depended critically on the subpopulations of juveniles that utilized coastal embayments (bays, lagoons, and estuaries). Conversely, the juvenile subpopulation along the exposed coast contributed negatively to overall population growth (lambda < 1) in 3 of the 4 years we conducted this study, due to elevated local mortality in that habitat. Life table response experiments confirmed that juvenile growth and survivorship were responsible for differences in lambda, and that nursery habitat choice could be a key contributor toward overall population fitness. Considering nurseries in a demographic source-sink context could aid conservation efforts by allowing identification or prioritization of the juvenile habitats most critical for population persistence.

Fodrie, FJ, Levin LA, Rathburn AE.  2009.  High densities and depth-associated changes of epibenthic megafauna along the Aleutian margin from 2000-4200 m. Journal of the Marine Biological Association of the United Kingdom. 89:1517-1527.   10.1017/s0025315409000903   AbstractWebsite

The Aleutian margin is a dynamic environment underlying a productive coastal ocean and subject to frequent tectonic disturbance. In July 2004, We used over 500 individual bottom images from towed camera transects to investigate patterns of epibenthic megafaunal density and community composition on the contiguous Aleutian margin (53 degrees N 163 degrees W) at depths of 2000 m, 3200 m and 4200 M. We also examined the influence of vertical isolation on the megafaunal assemblage across a topographic rise at 3200 m, located 30 km from the main margin and elevated 800 m above the surrounding seafloor. In comparison to previous reports from bathyal and abyssal depths, megafaunal densities along the Aleutian margin were remarkably high, averaging 5.38 +/- 0.43 (mean +/- 1 standard error), 0.32 +/- 0.02 to 0.43 +/- 0.03 and 0.27 +/- 0.01 individuals m(-2) at 2000 m, 3200 m and 4200 m, respectively. Diversity at 2000 M Was elevated by 15-30% over the deeper sites (3200-4200 m) depending on the metric, while evenness was depressed by similar to 10%. Levels of richness and evenness were similar among the three deeper sites. Echinoderms were the most abundant phylum at each depth; ophiuroids accounted for 89% of individuals in photographs at 2000 m, echinoids were dominant at 3200 M (39%), and holothurians dominated at 4200 m (47%). We observed a 26% reduction in megafaunal density across the summit of the topographic rise relative to that documented on the continental slope at the same depth. However, the two communities at 3200 m were very similar in composition. Together, these data support the modified 'archibenthal zone of transition' framework for slope community patterns with distinct communities along the middle and lower slope (the upper slope was not evaluated here). This study fills a geographical gap by providing baseline information for a relatively pristine, high-latitude, deep-sea benthic ecosystem. As pressures grow for drilling, fishing and mining on high-latitude margins, such data can serve as a reference point for much-needed studies on the ecology, long-term dynamics, and anthropogenically induced change of these habitats.

Fornes, WL, Demaster DJ, Levin LA, Blair NE.  1999.  Bioturbation and particle transport in Carolina slope sediments: A radiochemical approach. Journal of Marine Research. 57:335-355.   10.1357/002224099321618245   AbstractWebsite

In situ tracer experiments investigated short-term sediment mixing processes at two Carolina continental margin sites (water depth = 850 m) characterized by different organic C fluxes, (234)Th mixing coefficients (D(b)) and benthic assemblages. Phytoplankton, slope sediment, and sand-sized glass beads tagged with (210)Pb, (113)Sn, and (228)Th, respectively, were placed via submersible at the sediment-water interface at both field sites (Site I off Cape Fear, and Site m off Cape Hatteras). Experimental plots were sampled at 0, 1.5 days, and 90 days after tracer emplacement to examine short-term, vertical transport. Both sites are initially dominated by nonlocal mixing. Transport to the bottom of the surface mixed layer at both sites occurs more rapidly than (234)Th-based D(b) values predict; after 1.5 days, tagged particles were observed 5 cm below the sediment-water interface at Site I and 12 cm below at Site III. Impulse tracer profiles after 90 days at Site m exhibit primarily diffusive distributions, most likely due to a large number of random, nonlocal mixing events. The D(b) values determined from 90-day particle tagging experiments are comparable to those obtained from naturally occurring (234)Th profiles (similar to 100-day time scales) from nearby locations. The agreement between impulse tracer mixing coefficients and steady-state natural tracer mixing coefficients suggests that the diffusive analogue for bioturbation on monthly time scales is a realistic and useful approach. Tracer profiles from both sites exhibit some degree of particle selective mixing, but the preferential transport of the more labile carbon containing particles only occurred 30% of the time. Consequently, variations in the extent to which age-dependent mixing occurs in marine sediments may depend on factors such as faunal assemblage and organic carbon flux.

Frieder, CA, Nam SH, Martz TR, Levin LA.  2012.  High temporal and spatial variability of dissolved oxygen and pH in a nearshore California kelp forest. Biogeosciences. 9:3917-3930. AbstractWebsite

Predicting consequences of ocean deoxygenation and ocean acidification for nearshore marine ecosystems requires baseline dissolved oxygen (DO) and carbonate chemistry data that are both high-frequency and high-quality. Such data allow accurate assessment of environmental variability and present-day organism exposure regimes. In this study, scales of DO and pH variability were characterized over one year in a nearshore kelp forest ecosystem in the Southern California Bight. DO and pH were strongly, positively correlated, revealing that organisms on this upwelling shelf are not only exposed to low pH but also to low DO. The dominant scale of temporal DO and pH variability occurred on semi-diurnal, diurnal and event (days-weeks) time scales. Daily ranges in DO and pH at 7 m water depth (13 mab) could be as large as 220 mu mol kg(-1) and 0.36 units, respectively. Sources of pH and DO variation include photosynthesis within the kelp forest ecosystem, which can elevate DO and pH by up to 60 mu mol kg(-1) and 0.1 units over one week following the intrusion of high-density, nutrient-rich water. Accordingly, highly productive macrophyte-based ecosystems could serve as deoxygenation and acidification refugia by acting to elevate DO and pH relative to surrounding waters. DO and pH exhibited greater spatial variation over a 10 m increase in water depth (from 7 to 17 m) than along a 5 km stretch of shelf in a cross-shore or alongshore direction. Over a three-month time period, mean DO and pH at 17 m water depth were 168 mu mol kg(-1) and 7.87, respectively. These values represent a 35% decrease in mean DO and 37% increase in [H+] relative to near-surface waters. High-frequency variation was also reduced at depth. The mean daily range in DO and pH was 39% and 37% less, respectively, at 17m water depth relative to 7 m. As a consequence, the exposure history of an organism is largely a function of its depth of occurrence within the kelp forest. With knowledge of local alkalinity conditions and high-frequency temperature, salinity, and pH data, we estimated pCO(2) and calcium carbonate saturation states with respect to calcite and aragonite (Omega(calc) and Omega(arag)) for the La Jolla kelp forest at 7 m and 17 m water depth. pCO(2) ranged from 246 to 1016 mu atm, Omega(calc) was always supersaturated, and Omega(arag) was undersaturated at the beginning of March for five days when pH was less than 7.75 and DO was less than 115 mu mol kg(-1). These findings raise the possibility that the benthic communities along eastern boundary current systems are currently acclimatized and adapted to natural, variable, and low DO and pH. Still, future exposure of coastal California populations to even lower DO and pH may increase as upwelling intensifies and hypoxic boundaries shoal, compressing habitats and challenging the physiological capacity of intolerant species.

Frieder, CA, Gonzalez JP, Bockmon EE, Navarro MO, Levin LA.  2014.  Can variable pH and low oxygen moderate ocean acidification outcomes for mussel larvae? Global Change Biology. 20:754-764.   10.1111/gcb.12485   AbstractWebsite

Natural variation and changing climate in coastal oceans subject meroplanktonic organisms to broad ranges of pH and oxygen ([O2 ]) levels. In controlled-laboratory experiments we explored the interactive effects of pH, [O2 ], and semidiurnal pH fluctuations on the survivorship, development, and size of early life stages of two mytilid mussels, Mytilus californianus and M. galloprovincialis. Survivorship of larvae was unaffected by low pH, low [O2 ], or semidiurnal fluctuations for both mytilid species. Low pH (<7.6) resulted in delayed transition from the trochophore to veliger stage, but this effect of low pH was absent when incorporating semidiurnal fluctuations in both species. Also at low pH, larval shells were smaller and had greater variance; this effect was absent when semidiurnal fluctuations of 0.3 units were incorporated at low pH for M. galloprovincialis but not for M. californianus. Low [O2 ] in combination with low pH had no effect on larval development and size, indicating that early life stages of mytilid mussels are largely tolerant to a broad range of [O2 ] reflective of their environment (80-260 ╬╝mol kg(-1) ). The role of pH variability should be recognized as an important feature in coastal oceans that has the capacity to modulate the effects of ocean acidification on biological responses.

Frieder, CA, Gonzalez JP, Levin LA.  2014.  Uranium in larval shells as a barometer of molluscan ocean acidification exposure. Environmental Science & Technology. 48:6401-6408.   10.1021/es500514j   AbstractWebsite

As the ocean undergoes acidification, marine organisms will become increasingly exposed to reduced pH, yet variability in many coastal settings complicates our ability to accurately estimate pH exposure for those organisms that are difficult to track. Here we present shell-based geochemical proxies that reflect pH exposure from laboratory and field settings in larvae of the mussels Mytilus californianus and M. galloprovincialis. Laboratory-based proxies were generated from shells precipitated at pH 7.51 to 8.04. U/Ca, Sr/Ca, and multielemental signatures represented as principal components varied with pH for both species. Of these, U/Ca was the best predictor of pH and did not vary with larval size, with semidiumal pH fluctuations, or with oxygen concentration. Field applications of U/Ca were tested with mussel larvae reared in situ at both known and unknown conditions. Larval shells precipitated in a region of greater upwelling had higher U/Ca, and these U/Ca values corresponded well with the laboratory-derived U/Ca-pH proxy. Retention of the larval shell after settlement in molluscs allows use of this geochemical proxy to assess ocean acidification effects on marine populations.