CALIFORNIA SEA LIONS: an INDICATOR for INTEGRATED ECOSYSTEM ASSESSMENT Calcofi Rep., Vol

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CALIFORNIA SEA LIONS: an INDICATOR for INTEGRATED ECOSYSTEM ASSESSMENT Calcofi Rep., Vol MELIN ET AL.: CALIFORNIA SEA LIONS: AN INDICATOR FOR INTEGRATED ECOSYSTEM ASSESSMENT CalCOFI Rep., Vol. 53, 2012 CALIFORNIA SEA LIONS: AN INDICATOR FOR INTEGRATED ECOSYSTEM ASSESSMENT OF THE CALIFORNIA CURRENT SYSTEM SHARON R. MELIN, ANTHONY J. ORR, JEFFREY D. HARRIS, JEFFREY L. LAAKE, AND ROBERT L. DELONG National Oceanic and Atmospheric Administration National Marine Fisheries Service Alaska Fisheries Science Center National Marine Mammal Laboratory Seattle, WA 98115 ABSTRACT periods of large- and small-scale environmental changes, We examined the annual number of pups born, pup be sensitive to changes in the ecosystem, and have traits mortality, and pup weights of California sea lions (Zalo- that respond to and that can be measured in relation to phus californianus) at San Miguel Island, California, and the ecosystem processes of interest (Rice and Rochet related them to large and small-scale oceanographic 2005). Upper trophic level marine predators often make indices in the central California Current System (CCS) good indicator species because annual changes in popu- between 1997 and 2011. Annual variability in the num- lation parameters, such as births, mortality, and growth, ber of pups born and pup mortality was best explained are often linked to oceanographic changes (e.g., pro- by the mutlitvariate ENSO index (MEI) that tracks the duction of chlorophyll and zooplankton) that affect the El Niño/La Niña cycle. Annual variability in average pup distribution and availability of lower trophic level prey weights was best explained by a sea surface temperature (e.g., euphausiids, fishes, cephalopods) (Ainley et al. 2005; anomaly index (SSTA); average pup weights were lower Beauplet et al. 2005; Foracada et al. 2005; Reid and For- in years when the SSTA was greater than 1˚C above cada 2005; Reid et al. 2005; Wells et al. 2008). normal. We demonstrated that California sea lions are California sea lions (Zalophus californianus) are upper sensitive to large and small-scale changes in ocean condi- trophic level marine predators that are abundant and tions through changes in their reproductive success, pup permanent residents of the CCS. Their range extends growth, and pup mortality. Therefore, California sea lions from northern Mexico to Canada and much of their are an ideal indicator species for the IEA of the CCS. life history has evolved to take advantage of the high ocean productivity in the CCS. Weaning and reproduc- INTRODUCTION tion occur during late spring and early summer, respec- Integrated ecosystem assessment (IEA) is the scien- tively, during the peak upwelling period in the CCS tific foundation that supports ecosystem-based manage- when primary productivity is at its maximum (Bograd et ment (Levin et al. 2009). A central component of IEA al. 2009). California sea lion females give birth to a single is the identification of indicator species that respond pup between May and June that they provision through to changes in the ecosystem. In the California Cur- lactation. Lactation usually lasts 11 months during which rent System (CCS), large-scale global processes like the time females are central-place foragers, alternating 2–5 Pacific Decadal Oscillation (PDO), North Pacific Gyre day foraging trips to sea with 1–2 day nursing visits to Oscillation (NPGO), and El Niño Southern Oscillation the colony (Melin et al. 2000). Lactating females exploit (ENSO) as well as small-scale processes like localized the continental shelf, slope, and offshore regions of the disruption of seasonal upwelling can alter the trophic central and southern CCS throughout the year (Kuhn dynamics on time scales of months, years, or decades 2006; Melin et al. 2008), making more than 60 foraging (Hayward 1997; McGowan et al. 2003; Goericke et al. trips between the California Channel Islands and Mon- 2007; Bjorkstedt et al. 2010; King et al. 2011). In the terey Bay, California. Their large foraging area and diving CCS, the atmospheric forcing associated with the PDO capabilities give them access to a diverse prey assemblage and NPGO controls decadal patterns in upwelling and resulting in a diet that includes over 30 taxa of fish and results in regionally variable coastal upwelling condi- cephalopods (Antonelis et al. 1990; Lowry et al. 1990; tions that affect primary and secondary marine produc- Melin et al. 2010). tivity and consequently, the distribution of fishes and Over the past 40 years, population parameters of Cali- other higher trophic level marine organisms. Thus, suit- fornia sea lions have shown annual variability associated able indicator species for the CCS must be sensitive to with large- and small-scale oceanographic events. The marine ecosystem changes at various spatial and tempo- populations breeding in the California Channel Islands ral scales. An indicator species should be directly observ- off the southern coast of California experienced signif- able, have a historical time series of data that includes icant declines in births, increased pup mortality, lower 140 MELIN r3 lo.indd 140 10/30/12 9:06 PM MELIN ET AL.: CALIFORNIA SEA LIONS: AN INDICATOR FOR INTEGRATED ECOSYSTEM ASSESSMENT CalCOFI Rep., Vol. 53, 2012 mean pup weights, and changes in the diet in response METHODS to the warm oceanographic conditions associated with the El Niño phase of ENSO events in 1982–83 (DeLong Oceanographic Indices et al. 1991), 1992–93 (DeLong and Melin 2000), and PDO, NPGO, and ENSO. The PDO signal is 1997–98 (Weise and Harvey 2008; Melin et al. 2010). strongest north of 38˚N, the NPGO is strongest south The population effects lasted for 1 to 4 years (Lowry of 38˚N (Di Lorenzo et al. 2008), and the ENSO sig- and Maravilla-Chavez 2005). Furthermore, Califor- nal varies depending on the strength of the event at the nia sea lions are also sensitive to regional and localized equator (King et al. 2011) but all three indices are related changes in their foraging environment that affect their and affect the CCS (King et al. 2011). So, we explored prey base (Weise et al. 2006; Weise and Harvey 2008). In relationships between these indices and California sea 2009, a brief collapse of the summer seasonal upwelling lion population parameters. For each year between along the central California coast (Bjorkstedt et al. 2010) 1997 and 2011, monthly values for the PDO (http:// resulted in an unprecedented level of California sea lion jisao.washington.edu/pdo/PDO.latest), NPGO (http:// pup mortality, a dramatic change in the adult female diet, www.o3d.org/npgo/npgo.php), and ENSO (multivar- and contributed to a reduction in the number of births iate ENSO index (MEI), http://www.esrl.noaa.gov/ in the following year (Melin et al. 2010). The impact of psd/enso/mei/table.html) were averaged for: 1) October these anomalous oceanographic events on the California through the following June (average gestation period) for sea lion population are presumably meditated through models explaining trends in pup births, 2) June and July their affect on sea lion prey availability (i.e., distribution, for models evaluating pup mortality up to 5 weeks of abundance), but it is difficult to measure prey availabil- age, and 3) June through September for models explor- ity directly. Therefore indices of ocean conditions like ing variability in pup mortality and pup weights at 14 the PDO, NPGO, upwelling, and sea surface tempera- weeks of age. Because the MEI is measured at the equa- tures that affect distribution and abundance of prey can tor, the index was lagged 3 months, after testing lags of be used as proxies for prey availability to California sea 0 to 6 months, based on the highest positive correlation lions and consequently, may explain annual variability in between the average MEI values and local sea surface California sea lion population indices. temperatures in lactating female sea lion foraging areas A reduction in prey available to lactating California in the CCS (e.g., MEI value at the equator in January sea lion females has the greatest population effect because was assigned the CCS MEI value in April). it affects reproduction and survival of pups. When prey is Local Upwelling and Sea Surface Temperature. Large- scarce, lactating females expend more energy to meet the scale oceanographic patterns affect local ocean con- demands of reproduction by foraging farther from the ditions in the CCS through changes in the timing, colony and/or diving deeper presumably in response to strength, and characteristics of upwelling and changes changes in the spatial distribution of their prey (Melin et in sea surface temperature that affect the distribution of al. 2008). More importantly, movement of prey outside sea lion prey over shorter time periods (i.e., weeks or the normal adult female foraging range results in lon- months) and may have more immediate effects on Cali- ger foraging trips (Melin et al. 2008),which may result fornia sea lion population indices than large-scale pro- in slower growth or starvation of the pup if the for- cesses. We used upwelling and sea surface temperature aging trip durations exceed the pup’s fasting capability. indices to investigate the effect of small-scale ocean- In addition, because lactating females are usually also ographic conditions on California sea lion population pregnant during nine months of the 11-month lacta- and diet indices. The monthly coastal upwelling index tion period, a diet that is insufficient to support both at 33˚N 119˚W (UWI33) and 36˚N 122˚W (UWI36) lactation and gestation may result in the resorption of (http://www.pfeg.noaa.gov/products/PFEL/modeled/ the fetus or a premature birth. Given the relationships indices/upwelling/NA/data_download.html) between between ocean conditions, prey availability, and Califor- 1997 and 2011 was used as an index of regional monthly nia sea lion behavior, we used regional and local ocean- ocean productivity along the central California coast ographic and adult female diet indices as explanatory (Schwing et al.
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