Physical, biological and economic interconnections in the ecosystems and fisheries off California, 1877–2004

Quaternary International 310 (2013) 7e19

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Physical, biological and economic interconnections in the ecosystems and fisheries off California, 1877e2004 Jerrold G. Norton a, *, Janet E. Mason a, *, Cindy Bessey b, Samuel F. Herrick c a

Southwest Fisheries Science Center, NOAA/National Marine Fisheries Service, 1352 Lighthouse Ave., Pacific Grove, CA 93950, USA Florida International University, Miami, FL 33265, USA c Southwest Fisheries Science Center, La Jolla, CA 92037, USA b

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 16 November 2011

Comparisons of variation in physical, biological and economic indicators from 1877 to 2004 indicate that events in the physical ocean environment off California can be traced through the ecosystem to the commercial fishers’ economic environment. The annual landed weight of 29 commercial fish and invertebrate species, with various life histories and trophic associations, were used in an empirical orthogonal function analyses to isolate two major patterns of variation in the landings and form a graphical ecospace. This ecospace shows that dominant species associations have occurred in the landings during particular time intervals. These changes in ecospace are closely correlated to variation in physical environmental indices (r2 > 0.8). The analysis indicates that 50% of the landings variability is summarized in the two patterns, which represent variation at intervals greater than 40 years and less than 20 years. When the independently derived fish landings series are lengthened using published data and proxies, the physical to ecological to economic linkages are shown to occur from the late 19th century until 2004. When the purse seine fishery failure of the early 1920s and the growth of sardine cannery infrastructure from 1900 to 1945 are examined, results are consistent with perturbations beginning in the physical environment that are transferred through the ecosystem to influence the fishers’ harvest opportunities. These harvests will then influence future fisheries investment. Ó 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

1.1. Development and implications of an ecological space (ecospace)

Variations in the physical environment have been found to influence ocean species distribution and reproductive success (Parrish et al., 1981; Chelton et al., 1982; Beamish et al., 1999; Hollowed et al., 2007) at periods from less than a year to more than 1000 years (McGowan, 1990; Finney et al., 2002; Field et al., 2009). The variable physical environment has also been linked to intermittent success of commercial fisheries (Uber and MacCall, 1992; Mantua et al., 1997; Klyashtorin, 2001) and there is evidence that customs associated with northwest Native American utilization of anadromous fish were adapted to the effects of environmental variation (Johnsen, 2009). This report continues and augments studies linking the physical environment to the abundance of 29 commercially harvested species through intervals exceeding 75 years.

In previous studies, empirical orthogonal function (EOF) analyses of commercial landings records for 29 fish and invertebrates species showed the two functions or patterns (EOF1 and EOF2) with the most explanatory ability accounted for 54% of the temporal variability of these species during 1928e2004. These analyses detect recurring patterns in the landings time series and rate these patterns as a percentage of the overall variability of the landings data set. Landings were from an ocean area reaching 600 km off the California coast (Fig. 1). The low noise levels indicated by the large percentage of variance explained by EOF1 and EOF2, suggest that consistent landing of these 29 indicator species (Table 1) and the opportunistic commercial fishery contribute to the utility of the EOFs as ecosystem indicators (Norton and Mason, 2003; 2004; Norton et al., 2009). The indicator species are conspicuous members of benthic, coastal and pelagic habitats and have varied trophic and fishery associations within these habitats. When EOF1 and EOF2 are combined in a single diagram, with each species having a unique position, they indicate an ecological climate signal expressed as changing species

* Corresponding authors. E-mail addresses: [email protected] (J.G. Norton), Janet.Mason@noaa. gov (J.E. Mason), cbessey@fiu.edu (C. Bessey), [email protected] (S.F. Herrick). 1040-6182/$ e see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.10.041

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Fig. 1. Diagrams of the coast of North and South America between 10
S and 45
N (upper, right) and the coast of California from 30.5
N to 42
N (left). On the right, equatorial Kelvin waves travel across the equatorial Pacific ocean form west to east (dotted, open arrow). When they reach the coast of South America they are reflected north and south along the coast (small open arrows) as coastal Kelvin (planetary) waves that are partially trapped to the coast. These waves lose energy to slower moving Rossby waves, a second type of planetary wave, that propagate westward with phase speeds that are about ten times greater at the equator than at 30
N. The curved heavy, dashed line indicates the inverse relation of phase speed to latitude. Poleward currents near the coast (filled small arrows in both diagrams) are influenced by changes in the vertical density structure caused by planetary waves. Upwelling waves decrease northward flow and downwelling waves increase northward flow and warming. The NINO12 box (right) shows the ocean area were average sea surface temperature is measured for the OM1 index. The fish landings information used in this report was from the area within 600 km of the California coast (upper, right). Coastal geographical features of California (shaded) and major commercial fishing ports of the California coast are shown on the left. Small filled arrows represent near-shore, northward components of the California Current System and open arrows represent the broad, southerly flow of the California Current. Areas and distances listed on the upper right are approximate.

proportions in the total California landings. Fig. 2A is a diagram of changing species composition through the 1928e2004 interval as an ecological space or ecospace that reflects ecological processes affecting species abundance off California. The EOFs and the ecospace derived from them are robust to the removal of dominant species from the analyses, suggesting that the characteristics of EOF1 and EOF2 and their expression through the 1928e2004 interval, are shared by other major ecosystem components of the California Current System. The time series of EOF1 and EOF2 expression in individual years is highly correlated with time series of physical environmental indices computed on an annual basis (Norton and Mason, 2004; 2005). 1.2. Economic connections When species composition in the marine environment off California changes, harvest opportunities and associated economic opportunities change. Total landing weights for California commercial fishery declined from 1940 to 1965 as the species ensemble available to the fishers changed. This led to decreases in fishery infrastructure and investment. The number of boats making commercial landings in California decreased to about 2500 boats in 1965 from a maximum of about 6000 boats in the early 1940s. The percentage of boats longer than 21 m (65 feet) also decreased in this interval (Norton et al., 2009). The fishers’ behavior changed as species formerly harvested became less abundant and therefore more costly to harvest.

Given the strong global demand for seafood (O’Bannon, 2001; Herrick et al., 2009) the choice and quantities of species supplied will tend to reflect the relative abundance of commercially harvested species in the ecosystem. Therefore, the supply response on the part of fishers in the face of a persistent demand for the species that were consistently landed over the study interval would seem an effective way to characterize changes in the ecosystem in the absence of more complete biological information (Norton and Mason, 2004; Herrick et al., 2007; Swartz et al., 2010). 1.3. Progress reported here This report describes additional findings that lead in two related directions. First, the commercial fish and invertebrate landings and physical environmental indices used in earlier studies are extended to include earlier dates. Second, this paper seeks to confirm and expand understanding of apparent links from the physical to the ecological environments and from the ecosystem to fishers’ economic opportunities. Simple correlations test the hypothesized linkages. Three difficulties that lead to misinterpretation of correlation analyses are series autocorrelation (Chelton, 1983), chance occurrence of correlation in short series (Myers, 1998), and the interference of environmental cycles longer than the series analyzed. These problems are characteristic of most climate studies using historical data and they cannot be completely eliminated. However, extending the series length and adding variability that was not previously

J.G. Norton et al. / Quaternary International 310 (2013) 7e19

2. Regional setting

Table 1 Names of species used in the analysis. Albacore Anchovy (northern anchovy) Barracuda (Pacific barracuda) Bluefin tuna (Pacific bluefin tuna) Pacific bonito Butterfish (Pacific pompano) Cabezon California halibut Giant seabass Hake (Pacific hake) Herring (Pacific herring) Jack mackerel Lingcod Spiny lobster Dungeness (market) crab Ocean whitefish Pacific halibut Chub mackerel (Pacific mackerel) Sablefish Sardine (Pacific sardine) Scorpionfish (California scorpionfish) Sheephead (California sheephead) Skipjack (skipjack tuna) Market squid Swordfish White croaker White seabass Yellowfin tuna Yellowtail

9

o x

x o o x

x

ox

Thunnus alalunga Engraulis mordax Sphyraena argentea Thunnus orientalis Sarda chiliensis Peprilus simillimus Scorpaenichthys marmoratus Paralicthys californicus Stereolepis gigas Merluccius productus Clupea pallasii Trachurus symmetricus Ophiodon elongatus Panulirus interruptus Cancer magister Caulolatilus princeps Hippoglossus stenolepis Scomber japonicus Anoplopoma fimbria Sardinops sagax Scorpaena guttata Semicossyphus pulcher Katsuwonus pelamis Doryteuthis opalescens Xiphias gladius Genyonemus lineatus Atractoscion noblis Thunnus albacares Seriola dorsalis

oe not plotted in figures; xe not in 1920-2004 analysis.

in the analysis can reduce the likelihood and severity of these problems in correlation test interpretation. Proxy time series extend the correlated series into periods at the end of the 19th and beginning of the 20th centuries where physical to ecological to economic linkages have not been previously examined. In previous studies (Norton and Mason, 2004; Norton et al., 2009), apparently significant correlations were found between ecological variation as represented by time variation of the EOFs and a variety of remote and local physical indices. The EOF time variation shows significant co-variability to several large-scale indices, particularly those associated with the equatorial Pacific ocean. This co-variability suggests oceanic and atmospheric linkages to the ecosystem exploited by California fishers (Rasmusson and Wallace, 1983; Norton and McLain, 1994; Chelton and Schlax, 1996; Norton and Mason, 2005). The ecological space or ecospace, which gives a two dimensional representation of ecological changes as they evolve through time, are compared to the overall dockside value of the catch in Fig. 2A. This figure collects previously published data (Norton and Mason, 2005; Herrick et al., 2009; Norton et al., 2009) and combines it with new information that will be discussed in a qualitative economic sense. Two distinct and well-documented fishery economic events of the early 20th century are examined below. Pacific sardines have a unique role in the California Current ecosystem and in this study for at least three reasons. First, the total landings of sardines have been nearly equal to the landings of all other species during the 1916e2004 period (Marr, 1960; Murphy, 1966; Mason, 2004). Second, sardine variability is important in EOF1 temporal variability but not important in EOF2 temporal variability, which suggests that the time series of California sardine landings is similar to the time variation of EOF1 (Norton and Mason, 2003, 2004). Third, when viewed on multidecadal scales sardine biomass estimates and sardine landings time series have similar variability (Murphy, 1966; Herrick et al., 2007; Norton et al., 2009).

The coast of California extends from 32.5
N, 117.2
W northwest to 42.0
N, 124.2
W (Fig. 1). Commercial fishers landing their catch in California are influenced by three dynamic components of the California Current System. First, the California Current is an 800e1000 km wide meandering current that flows at rates of about 5e25 km/day southward along the west coast of North America between 48
N and 25
N (Fig. 1). When compared to the oceanic waters to the west the California Current has lower temperature and salinity and higher oxygen concentration, characteristics it carries from the subarctic Pacific (Reid et al., 1958; Lynn and Simpson, 1987; Di Lorenzo, 2003). Second, the inshore counter current is a generally northward flow within 150 km of the coast. At the surface the inshore counter current may have wind-forced reversals in spring and summer. The subsurface inshore counter current is more continuous in space and time and frequently has the strongest northward flow at the shelf break between 100 and 300 m depth. Often there is complex, banded poleward and equatorward flow in this nearshore region (Strub et al., 1987; Pierce et al., 2000). Third, a meandering, highly dynamic transition zone occurs 200e300 km offshore. This part of the southward flowing California Current is the transition between the region dominated by inshore processes and the southward flow of the eastern limb of the subtropical oceanic gyre (Lynn and Simpson, 1987; Parrish et al., 2000; Di Lorenzo, 2003). The California Current System is strongly forced by local and basin scale winds (Lynn and Simpson, 1987; Mantua et al., 1997; Parrish et al., 2000). There is also strong forcing within the ocean by locally and equatorially forced coastally trapped Kelvin planetary waves (Fig. 1). Planetary waves, which may lead to upwelling or downwelling of the ocean’s density structure, are observed to propagate northward along the coast at 30e90 km/day and radiate energy westward at 1e5 km/day as Rossby waves, a second type of planetary wave (Norton and McLain, 1994; Chelton and Schlax, 1996; Fu and Qiu, 2002). Ocean density structure changes induced by coastally trapped waves will change the intensity of the currents described above. These large-scale atmospheric and oceanic connections with the entire Pacific basin lead to Californian El Niño and La Niña events that are similar and generally in phase with those observed in the equatorial Pacific Ocean (Chelton et al., 1982; Rasmusson and Wallace, 1983; Chelton and Schlax, 1996). Equatorward winds associated with the eastern Pacific atmospheric high pressure system lead to spring and summer upwelling of deeper, cooler, higher nutrient water along much of the California coast (Bakun, 1990; Di Lorenzo, 2003). This coastal and offshore upwelling leads to photic layer nutrient enrichment and increased biological productivity in a coastal band that may be more than 500 km wide (Bakun, 1990; Rykaczewski and Checkley, 2008). Local wind-forced upwelling along the coast may be enhanced or attenuated by locally and remotely generated (Fig. 1) coastally trapped waves from the south. The majority of the California commercial fish and invertebrate landings (Mason, 2004) were made at or near the ports of Monterey, Los Angeles, and San Diego. Even though each species is weighted equally (Section 3.2), the analyses are biased toward central and southern California fisheries and ecosystems because more of the 29 indicator species are landed from this area. Los Angeles and San Diego also have extensive landings from the ocean south of California. These are excluded from the analyses because the origin of these harvests is less well known. The Southern California Bight, south of Point Conception, has variable bottom contours, basins and seven islands (filled in

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Fig. 2. Diagrams derived from California commercial fish landings EOF analysis. Panel A (above) shows an ecological space (ecospace) formed by plotting loading values for each species in EOF1, horizontal, and in EOF2, vertical axes. The dates give temporal scale. Species that have near maxima in the same period are looped together. As the ocean environment has changed, species associations have changed, as shown by clockwise rotation (gray arrow). The position in ecospace indicates the state of the California Current ecosystem. Four species with low loading values are not shown (Table 1). The small tables under each date show the 5-year means of commercial fishing statistics and are given at 5- and 10-year intervals to show maxima and minima. The entries from the top of the tables give: 1) total landings value, in millions of year-2000 US dollars ($M), and the percent of this total from the dockside sale of the 29 indicator species, 2) the value and the percent of total value of the combined catch of herring, anchovy, and sardines and 3) the total landings in millions of metric tons (Mmt) and the percent of this total from the landings of the 29 indicator species. Parts of the material summarized in Fig. 2 appear in Norton and Mason (2005), Herrick et al. (2009) and Norton et al. (2009). Panel B gives a second representation of the changing ecological state as time variable coefficients (TVC) of EOF1 (horizontal) and EOF2 (vertical). TVCs give the weight of their respective EOFs in each year from 1928 to 2004 (solid line). Start (S), end (E) and selected years are marked. The dotted line gives a two dimensional environmental index, with ocean mode one (OM1) on the horizontal and ocean mode two (OM2) on the vertical. All series are standardized for comparison. The square of the correlation coefficient (r2) value on the horizontal is 0.85 for the TVC1/OM1 correlation. On the vertical scale, the r2 value is 0.81 for the TVC2/OM2 correlation. See the text for additional details.

Fig. 1). The Santa Barbara Basin, between the mainland and three of the northern islands, at 34.2
N, 120.0
W, is about 590 m deep at the center (Barron et al., 2010). Its suboxic waters (Sigman et al., 2003) and annually varved, suboxic sediments preserve chronologies of fish scale deposition reaching hundreds of years into the past (Soutar and Isaacs, 1974; Berger et al., 2004; Field et al., 2009).

3. Methods This report uses information from a variety of independent sources and, where possible, develops time series to be compared by simple correlation. Partial review of material reported elsewhere provides contexts for the development of the more detailed, longer series presented below.

J.G. Norton et al. / Quaternary International 310 (2013) 7e19

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Fig. 3. The EOF analysis is extended to 85 years from 1920 to 2004. In panel A TVC1 (solid) is compared to OM1 (cross, dotted), where the r2 is 0.90. In panel B TVC2 (broken, heavy) is compared to OM2 (triangle, dash), where, r2is 0.83 with a trend correction to OM2 of 0.015 per year. The pre-correction r2 is 0.71 (see Section 3.4 for details). All series are standardized and may be offset for visual comparison.

3.1. California commercial fish and invertebrate landings The empirical orthogonal function (EOF) or principal component analysis depends on the continuous landings records of 29 species of fish and invertebrates (Table 1). These records were compiled by the California Department of Fish and Game and published in their Fish Bulletins (http://libraries.ucsd.edu/apps/ ceo/fishbull/index.html) from landing receipts listing the weight of each species and the amount paid to each boat at dockside for the weight of that species.

The Environmental Research Division of the Southwest Fisheries Science Center (Mason, 2004) has converted these landings weight tabulations into a database (CACom), which is available at http://www.pfeg.noaa.gov/products/las.html. Annual total landings from CACom are used in conjunction with annual landings information for the 1916e1927 period, published in California Department of Fish and Game Fish Bulletin number 74. The 29 indicator species have contributed from 64% to 98% of the annual total weight and from 64% to 71% of the annual total value of the landings of fish and invertebrates caught within the area from the

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California coast west to 600 km, and landed in California ports (Figs. 1 and 2A). Commercial landing tabulations do not represent rigorous scientific sampling, but they are more extensive in time, space and magnitude than any existing scientific samples from the California Current ecosystem. Consistent landings of the 29 species, through the study interval, indicate a dependable market demand. This suggests that markets will buy fish when they become available and fishers will supply fish as long as the costs of harvest do not exceed the revenue received for their catch. Demand appears to exceed domestic supply in most US seafood markets including California (O’Bannon, 2001; Swartz et al., 2010). Human population increases in California and around the globe have provided increasing markets for California fish and invertebrates. From 1930 to 2000 the human population of California increased six-fold. During the same period, annual per capita consumption of edible fish in the United States grew from 4.5 kg in 1930 to 7.1 kg in 2000. At the same time, the US population has more than doubled from 1930 to 2000 leading to a 340% increase in US fish consumption (O’Bannon, 2001; Swartz et al., 2010). The threefold increase in the world’s population and foreign market increases through the 20th century have created continuing harvest incentives for California commercial fishers (Swartz et al., 2010). These market and demographic statistics suggest that demand from domestic and foreign markets puts continuous harvest pressure on the 29 indicator species and on California’s ocean ecosystem.

3.2. Empirical orthogonal function (EOF) analysis Computing empirical orthogonal functions (EOFs) uses standard methods (Kutzbach, 1967; North et al., 1982), following the conventions of Norton and Mason (2003, 2004). These are reviewed briefly. Let the data matrix, [D0 ], from CACom consist of species with annual catch in rows corresponding to years. All columns are natural log transformed,

½DL  ¼ loge ½D0:

(1)

Standardizing [DL] by subtracting the mean of each column and dividing by the standard deviation of that column gives [D]. Derive the correlation matrix, [R], of [D] by

½R ¼ k½DT ½D:

(2) T

where k depends on the dimensions of [D] and [D] is the transpose of [D]. Then the eigenvectors [E] and Eigenvalues are derived from,

½R½E ¼ ½L½E:

(3)

The diagonal elements of [L] are eigenvalues (ln) that correspond to the column eigenvectors of [E]. EOF loadings (one value for each of n columns or species in [D0 ]) are given by

EOFn ¼ ðln Þ1=2 ½En 

(4)

where, ln gives the variance explained by EOFn. The first EOF, EOF1, has the largest ln and explains the most variance in [D]. EOF2 has the second largest ln, and so forth. The method of North et al. (1982) is used to find that EOF1 and EOF2 are significant below the 0.05 probability level (p < 0.05). The time variation of EOFn over the sampling interval (time varying coefficients) is given by

½TVCn  ¼ ½D½EOFn 

(5)

The EOFs and their TVCs, that give the variation of the EOF through time, are the ecological indicators. They are empirical in

that they are derived from data [D] and they are ecological in that they represent more than 20 inter-related species. The following analyses focus on two data matrices. The results from a 29 indicator species matrix covering 77 years (1928e2004) from Norton et al. (2009) and from a 24 species subset of the 29 species (Table 1), extended for this study to cover 85 years from 1920 to 2004 (Fig. 3). Although usable data for some species begins in 1916, it is not complete enough to extend the EOF analysis to dates earlier than 1920. Loading values for EOF1 and EOF2, which represents average states derived from the transformed data matrix, [D], are plotted as abscissa and ordinate, respectively, in Fig. 2A. These define the ecospace or ecological space that is an important part of this report. 3.3. Sardine abundance series: 1848e2004 Throughout the EOF studies, the sardine series was found to be strongly loaded in EOF1 and weakly loaded in EOF2 (Norton and Mason, 2004, 2005; Norton et al., 2009). Therefore, in the absence of complete records of the 29 indicator species, sardine abundance estimates are used as an EOF1-proxy to extend present analyses into the late 19th century. This is possible because suboxic waters (Sigman et al., 2003) at the bottom of the Santa Barbara Basin have preserved varved chronological records of sardine scale deposition rates (SDR). The scales are found within annual laminations beneath a portion of the sardine spawning grounds in the Southern California Bight (Fig. 1). The near absence of whole fish in the sediments suggests that the scales are deposited as the result of passive or forced shedding from live fish (Soutar and Isaacs, 1974; Baumgartner et al., 1992; Field et al., 2009). The varve ages that were used in forming chronologies have a temporal error of 3years (Schimmelmann et al., 1990; Berger et al., 2004; Field et al., 2009). For the analysis, a necessary simplification is to take sardine scale deposition rates as measurements of sardine abundance over the population range. Sardine landings and biomass (weight of the sardine population) estimates follow similar multi-decadal patterns, but the biomass estimates are derived from a variety of sources and estimate abundance independently of the fishery. The SDR estimates are not directly fishery dependent before about 1920 (Soutar and Isaacs, 1974). Therefore, sardine biomass estimates were used rather than landings in forming the combined 1849e2004 time series of sardine abundance. In an effort to maximize the SDR information presented in published studies, the five-year SDR estimates of Soutar and Isaacs (1974) and the 10-year estimates of Baumgartner et al. (1992) were standardized and averaged. Fortunately, the five- and 10year SDR estimates were comparable, except for temporal resolution. The SDR record was then smoothed with a three point running mean (Field et al., 2009) to make the series more comparable with recent biomass estimates. Recent biomass measurements (Hill et al., 2006) were adjusted to the same range as the SDRs, using linear correlation during the 1932e1955 common period, where r2 ¼ 0.76. The natural log transform makes the EOF1-proxy (SDR) analyses and the EOF analysis comparable over the 1877e2004 period. The natural log transform and standardization (Equations (1) and (2)) of all fish series restrains the determining effects of large sardine landings in the analysis (Norton and Mason, 2005). Therefore, the composite sardine series is a reasonable EOF1-proxy. 3.4. Physical environmental indices (ocean modes) Over the course of these studies, relevant to California commercial landings, the co-variation of several physical variables

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with the TVCs have been explored, including local and remote wind and SST variables and the Pacific Decadal Oscillation (PDO). These physical variables were examined in terms of: (1) apparent explanatory power (more than 60% of TVC variance), (2) homogeneous series length and (3) relevancy in terms of known physical processes. Common variability is found in all the large-scale indices examined (Norton and Mason, 2003, 2004; 2005; Herrick et al., 2007; Norton et al., 2009). The first source variable used in the present studies is the sea surface temperature (SST) anomaly from the Scripps Institution of Oceanography pier in La Jolla, California (32.9
N, 117.3
W), near San Diego (http://shorestation.ucsd.edu/active/index_active.html% 23lajollastation), abbreviated LSST below. The LSST has been discussed extensively (Marr, 1960; Jacobson and MacCall, 1995; McGowan et al., 1998; Norton, 1999) and is used in management of the sardine fishery (Hill et al., 2006). The second source variable is the annual average sea surface temperature anomaly series from an area at the extreme eastern side of the equatorial Pacific Ocean, 0
e10
S, 80
e90
W abbreviated Nino12 in the following (http://iridl.ldeo.columbia.edu/SOURCES/.Indices/.nino/.EXTENDED/. NINO12/gridtable.tsv). Choice of the Nino12 source variable is based on: (1) known linkages between the California Current and the equatorial Pacific (Rasmusson and Wallace, 1983; Chelton and Schlax, 1996) and (2) its availability in a time series that begins in 1858 and extends to the present (Kaplan et al., 1998; Reynolds et al., 2002). The first step in developing the derived ocean process indices from these source variables is to form annual anomalies from the overall arithmetic mean. This gives them a time-step that is identical to the TVCs. Then, two approaches were used in developing the ocean modes of variability: (1) serial integration and (2) filtering with a tapered smoother. Anomalies from mean Nino12 series, X, were serially integrated through time,

aXðyÞ ¼

y X

XðiÞ;

(6)

i¼b

where aX(y) is the accumulation of the anomaly time series X, b is the first year of the accumulation and y is a year from b to 2004. The annual value of the serial integration of the Nino12 annual anomaly, is termed “ocean mode one” (OM1). The use of tropical Nino12 to form OM1, a physical environmental index for the California Current region, points to a biologically important physical oceanic connection by coastally trapped wave processes between the eastern equatorial Pacific and the California Current. In addition, accumulating the anomaly of the physical variable matches the natural log of the population size in populations that change according to the exponential of the physical anomaly (Norton et al., 2009). Annual mean sea surface temperatures taken at La Jolla (LSST) were developed into the ocean mode two (OM2) index. The LSST values are local to the fish’s environment where variably lagged life cycle responses occur over periods of one to five years temporally smoothing TVC2 variation. An eleven point, centered, geometrically stepped, running mean of LSST seemed best in accommodating this variability in fish and invertebrate life history. Experimental trends of j0.005j to j0.02j
C per year were added to OM1 and OM2 and the correlations calculated iteratively to find the slope with the highest correlation coefficient. This procedure addresses two problems of using anomalies for retrospective ecological studies, but unfortunately it does not separate them. First, this is similar to finding the most biologically sensitive mean value of OM1 and OM2 for the changing biological systems studied. The arithmetic mean, which has

13

adaptive attraction and might be more important in an unchanging system, is used as an initial estimate of the most biologically sensitive mean (Rose and Lauder, 1996). Second, this procedure has the effect of adjusting overall trends in OM1 and OM2 to trends in TVC1 and TVC2 that arise from cycles that cannot be resolved within the time-interval considered (McGowan, 1990). This procedure was only effective in improving the relationship between TVC2 and OM2 (Fig. 2B). Significance levels for all correlations were adjusted for effective degrees of freedom determined by the long-lag (20%e30%) correlation method (Chelton, 1983). Smoothing and integration produce autocorrelated series with degrees of freedom that are lower than the number of values. However, the series used in this report run from 77 to 128 years in length and have sufficient corrected degrees of freedom to produce nominally statistically significant results. In general, correlation coefficient magnitudes larger than 0.8 (r2>0.64) are likely to occur by chance at a rate less than one in 20 trials (p < 0.05). The variables, TVC1, TVC2, OM1 and OM2, were also tested for their correlation relationship to random series and series independently derived from the ocean environment. When uniform random series were reddened in a first order sense (Rudnick and Davis, 2003), sampled and correlated 10,000 times to the four variables, the r2 threshold for the p < 0.05 level was 0.64 for TVC1 and less for the other three variables. To extend the tests of correlation uniqueness to randomized series of ocean origin, randomly assembled temporal sections of annual sea temperature series derived from the analysis of Palmyra Island coral (Cobb et al., 2003) were examined. The probability of the four variables apparently correlating by chance at the p < 0.05 level in 10,000 tests was greater for each variable in this test system. The p < 0.05 threshold r2 values for TVC1 and OM1 were 0.69 and 0.64, respectively. Threshold values for the other two variables were less. This kind of a test has the advantage of using series that represent measured ocean variability. However, there is the difficulty of possible causality linking the correlated variables and elevating the apparent p < 0.05 threshold (Cobb et al., 2003; Norton and Mason, 2005). 3.5. Economic indicators The value of the landings (fishers’ revenue) was obtained from trip tickets completed by fish dealers as a record of the transfer of fish from the fisher into the shore-side economic system (ex-vessel or dockside value). These records were treated in the same way as records of weight landed (Mason, 2004) and are available from the California Department of Fish and Game Bulletins. Other time series indexing the fishers’ economic environment have been more difficult to assemble, but there are two welldocumented economic events examined in terms of physical environmental and ecosystem changes: (1) the southern California purse seine fishery failure of 1920e1925 and (2) the construction of canneries along the coast of central and southern California in the first decades of the 20th century. Skogsberg (1925) estimated that investment in purse seine boats in southern California during 1920 was about 20 million US dollars, inflation adjusted to year-2000 ($M). The multi-species southern California purse seine fishery began with one boat in 1893 and grew steadily to more than 100 boats in the fishery in 1920. Most of these boats offloaded their catch in the Los Angeles Harbor area, which includes San Pedro, Terminal Island, Wilmington, and Long Beach. The purse seine, which involves encircling a school of fish with a vertical net and then closing or pursing the net at the bottom, remains one of the most efficient ways of harvesting schooling fish of all sizes.

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In southern California, purse seine fishers sold bluefin tuna and yellowtail to Los Angeles Harbor canneries that had originally been built to can sardines and mackerel (Smith, 1895; Scofield, 1954; Uber and MacCall, 1992). The canneries had developed methods of canning albacore tuna, but the diving behavior of albacore makes them difficult to catch using the encircling purse seine. Barracuda and white seabass were sold primarily in the fresh-fish markets and made up about 25% of the purse seine landings (Skogsberg, 1925). The California sardine canneries were constructed along the California coasts between 1893 and 1950. The canneries represented multi-million dollar investments in California fisheries. Before the canning process was introduced to California fisheries in the mid-1800s, salting and drying were the most common fish preservation methods. Canning processes opened worldwide markets for highly nutritious California-caught seafood in a stable form. The first sardine canneries were in Los Angeles Harbor and San Diego (Smith, 1895; Uber and MacCall, 1992). As the canneries became increasingly important in California fishers’ economic environment, the Los Angeles Harbor and San Diego canneries diversified to include tuna and San Diego canneries later specialized in tuna products. Canneries used offal from canning and some whole fish in reduction processes that produced fish meal and oil. Reduction became a valuable part of the sardine business and the subject of considerable regulatory debate. Monterey area canneries specialized in sardines and other small pelagic species and received the greatest sardine landings from the mid-1920s through the mid-1940s when total west coast landings grew from 64,000 metric tons (mt) to 614,000 mt. During the following decline of the sardine resource, southern California received the greatest sardine landings. The first Monterey sardine cannery was completed in 1902 in the southern bight of Monterey Bay. By 1918 eight more canneries had been built in Monterey, suggesting success by the first cannery. By 1945, there were 21 canneries in Monterey (Scofield, 1954; Uber and MacCall, 1992). Canning and reduction processes expanded at Moss Landing, 20 km north of Monterey, where operations grew from one cannery in 1935 to five canneries in 1945. There was also growth in canning and reduction throughout California as 30 canning and reduction plants in 1930 grew to more than 87 in 1945 and 132 in 1950, but part of the state wide growth was due to expansion of the tuna canning industry (Lindner, 1930; Scofield, 1954). Until the mid-1920s, sardine landings in Monterey were taken from within the southern bight of Monterey Bay, often within sight of the canneries (Lindner, 1930). Sardine landings did not appear limited by resource availability until the mid-1920s (Lindner, 1930; Clark, 1939). As the California harvests of sardines increased from a five-year average of 43,000 mt in 1922 to an average of 155,000 mt in 1927 fishers had to travel farther from port to fill cannery orders. For the Los Angeles Harbor fishery 98% of the sardine harvest was taken in Santa Monica Bay and along the mainland in the early 1920s, but less than 30% was taken from these local fishing grounds at the end of the 1920s. In Monterey 95% of the sardine harvest was within 20 km of port in the early 1920s, but by the end of the decade, less than 40% of the harvest was taken form these waters. By the late 1920s boats from Monterey were often traveling more than 100 km from port to fill cannery orders (Lindner, 1930). 4. Results Temporal associations of species in the ecological space or ecospace and the overall economic consequences of these associations for the 1928e2004 period are presented in Fig. 2A. Then the ecospace analysis is extended in five ways: 1) the time variable

Fig. 4. Landings of seven fish species commercially harvested during 1916e1935. Panel 4A shows the variation in three species of fish delivered to southern California canneries. Filled symbols on the solid line give the variations in albacore annual landings. Lines without symbols give bluefin tuna (broken) and yellowtail (solid) annual landings. Variation in barracuda (symbol, solid) and white seabass (solid) landings for the fresh-fish markets is shown in Panel 4B. Variation in butterfish (symbol, broken) and anchovy (symbol, solid) are shown in Panel 4C.

coefficients (TVC) of EOF1 and EOF2 were compared to physical environmental indices derived from Nino12 SST anomaly (OM1) and southern California LSST anomaly (OM2) in Fig. 2B, 2) the EOF analysis was extended to the 1920e2004 period using a 24 species subset of the 29 indicator species (Fig. 3A,B), 3) landings time series for five indicator species were examined through the 1916e1935 interval (Fig. 4) to show how environmental fluctuation influenced the economics of the southern California purse seine fishery, 4) sardine biomass was used as a proxy for TVC1 and compared to OM1 for the 1877e2004 period (Fig. 5), and 5) the 1850e1916 period was examined in terms of the investment incentives offered by an estimated 9 million metric tons of sardine off the west coast of the US. Changes in the relative proportion of species in the California landings for the 1928e2004 period are shown in Fig. 2A, which was formed by plotting EOF1 and EOF2 loading values for the 29 indicator species. Species grouped together within a closed loop have landings maxima in the time periods indicated. Maxima are relative to the catch for that species and the weight landed for a particular species may be orders of magnitude different from other species in the group. The species maxima are not necessarily synchronous and may occur in a particular year or years or through a group of years extending over a decade. Consequently, periods of landings maxima do not progress regularly in time within the loop. Broken lines on the advancing edge of each loop indicate periods of ecosystem and fishery transition from one group to the next.

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Fig. 5. The broken line is the TVC1-proxy, the natural log of annual sardine biomass estimates derived from scale deposition rates (SDR) before 1932 and sampling based biomass estimates after 1932. The TVC1-proxy is compared to OM1 (solid, cross). The OM1 series starts in 1856 and begins to track the TVC1-proxy in 1877. The r2 is 0.82 for the 1877e2004 period. Series are standardized and may be offset for visual comparison.

4.1. Extending the ecospace relationship: 1920e2004 It is possible to extend the EOF analysis to include 1920 using 24 of the 29 indicator species. If the relationships obtained for 1928e2004 period are not weakened in the extended analysis, then it will support the hypothesis that these physical to ecosystem linkages hold through time. The percentage of variance explained by the first two EOFs is 49% for the extended analysis and 54% for the 1928e2005 analyses. The fit of OM1 to TVC1 is similar to previous results with r > 0.9 (Fig. 3A). It appears that OM2 is a reasonable indicator of three to 20-year ecological perturbations shown TVC2 series. Excursions representing local increases in SST and increases in TVC2-values occurred during the 1920e1929, 1952e1959, 1974e1982, and 1983e1997 time periods. Excursions to lower SST and lower TVC2values occurred in 1938e1950, 1960e1972 and 1998e2002. The well-documented 1974e1982 climate shift (Norton et al., 1985; Ebbesmeyer et al., 1991) is the largest OM2 and TVC2 excursion of the 1920e2004 interval and the 1960e1982 period defines an approximately 20 year cycle (Fig. 3B). This is a longer cycle than previously reported for TVC2 (Norton and Mason, 2004; Norton et al., 2009). 4.2. Fisheries and economic fluctuations: 1916e1935 Landings records for seven fish species during the 1916e1930 period are useful in examining the southern California purse seine fishery failure of the early 1920s (Fig. 4). In 1919 the purse seine landings of albacore, barracuda, bluefin tuna, white seabass and yellowtail were all relatively high, suggesting a transition period in ecospace when species from adjacent areas are available for harvest. Albacore, bluefin tuna and yellowtail were cannery species, but only bluefin tuna and yellowtail were accessible to purse seine fishers, because albacore usually dive out the bottom of encircling nets. Barracuda and white seabass were available to purse seiners. Anchovy is one of the most common albacore forage species, but anchovy and butterfish were not important in the purse seine fishery during 1916e1935.

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The character of CACom landings data when it is not log transformed, standardized and summarized by EOF analysis is shown in Fig. 4, but the generalities of ecospace are evident in these records. In 1920 yellowtail and bluefin tuna landings dropped and albacore landings increased, showing a shift from the right toward the lower left in ecospace. Barracuda landings also fell in 1920. In 1921 bluefin tuna landings fell to below 2000 metric tons and remained low until 1926. Yellowtail landings also fell, as albacore landings remained relatively high. White seabass landings in the purse seine and other fisheries declined from 1919 to 1924 (Skogsberg, 1925). These changes in landings imply a temporary shift from right to lower left in ecospace (Fig. 2A). This temporary shift is also seen in the landings of anchovy beginning in 1919 (Fig. 4C). Butterfish appeared to herald the shift in 1919. Comparison of these shifts in landings to TVC2 and OM2 (Fig. 3B) suggests that the physical environmental and ecological conditions that made target species available to southern California purse seiners were not stable on a short-term (TVC2) basis. Albacore landings, primarily from fisheries methods other than purse seining, were high and bluefin tuna landings low (lower left in ecospace), until the mid-1920s when albacore landings fell to low levels and more than 2000 metric tons of bluefin tuna were landed in five consecutive years (Fig. 4A). These observations are consistent with the rebound (towards center on the vertical axis in ecospace) of the TVC2 and OM2 environmental variables (Fig. 3B). This rebound is shown in landings of the relatively long-lived migratory species (Fig. 4A) and the smaller shorter-lived species (Fig. 4C). A similar perturbation is seen in the cyclic progression of variables during the 1955e1965 period (Fig. 2B), where there are relatively large vertical but modest horizontal changes. Both vertical and horizontal changes in ecospace will affect the costs of harvests and subsequent investments. The economic impacts of TVC2 changes may be intense and short lived while changes brought by TVC1 may be more difficult to detect, but longer lasting (Fig. 2A). The 1916e1926 ecosystem changes had severe economic consequences for purse seine fishers and for canners who did not anticipate large changes in available species. Skogsberg (1925) estimated that investment in boats during 1920 was about 20 $M (year-2000 dollars). In 1917 and 1918 a fisher’s single “share” for a seasonal fishery was as much as $20,000. Aboard some purse seine vessels a “share” exceeded $30,000 in 1919, suggesting relatively low expense and effort for relatively high catch rates. Total trip revenues were divided into 9 to 12 shares. In 1917, 1918, and 1919 there was incentive, based on recent success, for purse seine fishers to increase their harvest capacity. Boat owners borrowed from banks and canneries to upgrade their boats and equipment. However, the landings records (Fig. 4) and the environmental indices (Fig. 3B) show important ecosystem changes during the early 1920s that decreased chances of continued success. Decreased landings of target species and resulting economic hardships caused many of the purse seine boats that had come from northern ports to leave southern California. Of the 50 southern California purse seine boats that fished for bluefin tuna in 1922, only a few were successful. Economic losses from investment in boats and fishing gear were heavy and by 1925 only a few of the 50 purse seine boats remaining in southern California were not mortgaged to canneries or banks (Skogsberg, 1925). 4.3. Sardine biomass, TVC1-proxy and cannery investment The variation in sardine biomass may be used as a proxy for EOF1 variation, or TVC1, because sardine has high loading in EOF1 and negligible loading in EOF2 (Fig. 2A). These findings are combined with the availability of the SST anomaly time series at Nino12 (Kaplan et al., 1998) to form a 128-year comparison of OM1

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and the TVC1-proxy (Fig. 5). Initially, the OM1 index has to accumulate information (Equation (6)) before it begins to track the TVC1-proxy in about 1877. The initial coincidence of the two curves in 1877 is near an apparent minimum in TVC1-proxy and OM1 (Fig. 5). Overall, the OM1 index sometimes leads and sometimes follows the TVC1-proxy. The TVC1-proxy appears to lead the OM1 values at the 1900s maxima, but it is possible that because of temporal inconsistencies in the two independent series the maxima occur at nearly the same time. The OM1 and TVC1-proxy track well from the onset of modern, more inclusive, biomass estimates until the minimum in 1964. During the 1960e1985 period the biomass estimates that give the TVC1-proxy are based on sparse egg and larva sampling of the depleted resource (MacCall, 1979; Barnes et al., 1992). A more accurately determined biomass, if available, might track the OM1 index more closely during the 1967e1985 period. OM1 appears to lead the TVC1-proxy after 1985. The patterns of variation before 1880 in the SDR record suggest a sardine biomass maxima in the 1840e1860 period, then a sharp decrease in sardine biomass followed by an increase again to a maximum around 1900, in a 40e50 year cycle (Fig. 5). The modern cycle from 1900 to 1998 suggests a cycle of 90e100 years in sardine biomass and TVC1 variation. California sardine canneries were constructed from the period of maximum sardine abundance in the early 1900s, through the period of largest sardine landings in the 1930s and early 1940s and into the period of sardine population decline. In 1940 during the period of maximum California landings there were 18 canneries in the Monterey Bay area. This number more than doubled in the next ten years, even though the sardine populations were decreasing. It was probably the persistent climate, TVC1-nature of sardine resource change that led to over investment in canneries. Investments in canning and reduction of sardines followed similar patterns in the San Francisco and Los Angeles areas, however it is more difficult to assess the degree of over investment in southern California because of growth in the tuna canning industries concomitant with the failure of the sardine resource. During the 1920s and 1930s the sardine biomass in the coastal waters off the west coast was estimated to be three to four million metric tons (Murphy, 1966; MacCall, 1996). However, this analysis suggests that during the great sardine harvests the biomass was experiencing an environmentally induced decline from a peak of nine million metric tons in the early 1900s (Fig. 5). Intense harvests by an increasingly effective industrial fishery may have increased the rate of sardine population decline and affected other species of the California Current System’s ecosystem. 5. Discussion The relation of the physical environmental and economic variables to the ecospace diagram indicates linkages from the physical environment to the California fishers’ economic environment. Much work remains to show the details of the implied linkages. However, several characteristics of the co-varying systems support the validity of the proposed linkages and the generality of the results. The large number of species used in this analysis that extends over eight decades supports the ecological interpretation of the ecospace diagram (Myers, 1998). Commercial landing tabulations in CACom do not represent rigorous scientific sampling, but they are more extensive in time, space and magnitude than any available scientific samples. Even though the annual total landings for many CACom species are the result of hundreds to thousands of records, inaccuracies probably remain. Consequently, the signals detected by EOF1 and EOF2, which account for 50% of the overall variance, may represent even more of the population variance for many California marine ecosystem components.

Position changes in ecospace indicate changes in the fishers’ economic environment (Fig. 2A). When one species decreases in the landings and another species appears to replace it in the landings, as is indicated by a shift from one area to another in ecospace, it reflects changes in total weight landed, in harvest location, in harvesting gear, in the type and number of boats in the fishery. A large change in ecospace and total ex-vessel value occurred from 1945 to 1965 (Fig. 2A). This large and persistent change in physical, ecological and economic climate is shown in OM1 and TVC1 (Fig. 2A,B). A shorter, OM2 and TVC2, change affected the southern California purse seine fishery in the early 1920s (Figs. 4 and 3B). It is unlikely that fishers working in largely unregulated fisheries with expanding markets would have been vulnerable to these economic changes without ecosystem changes that limited harvests. 5.1. Time series Useful simplifications of complex data allow the use of linear approaches throughout the analyses. If it were possible to examine the entire ecosystem, then layered, non-additive responses of complex nature would certainly be encountered. Time series of different lengths may have different results in EOF analyses (Yuan et al., 1997). When the landings series is shortened to begin in the late 1940s or early 1950s, TVC1 is dominated by the 1973e1978 climate shift in a pattern similar to TVC2 and several concurrent California Current ecological time series (McGowan et al., 1998). The inclusion of earlier years in the analysis allows this climate shift to fall into EOF2, and shows the importance of longer time series in detecting lower frequency more energetic cycles that may be influencing ecosystems and the fisheries that depend on them. The longest time series available were compared, so that inferences have the greatest chance of generality (Chelton, 1983; Myers, 1998). Physical environmental indices (OM1, OM2), derived from Nino12 SST and LSST anomalies, respectively, are annual averages from daily and monthly values. Nino12 SST is also an area-average. The serial integration of Nino12 SST annual anomalies to give OM1 follows the integrative pattern of population growth, where the natural log of biomass grows as a function of the serial integration of the physical environmental anomaly (Norton et al., 2009). Population decline or growth (accumulation) over decadal periods is a major contributor to EOF1 ecosystem processes (Figs. 2 and 5). Physical and biological cycles between 40 and 170 years length have also been detected in single species analyses of physical and fisheries time series from eastern north Pacific and north Atlantic oceans (Beamish et al., 1999; Klyashtorin, 2001; Finney et al., 2002; Berger et al., 2004). 5.2. Ecosystem responses to cyclic environmental variability Many periods of ocean fluctuation are observed (McGowan, 1990; Finney et al., 2002; Hollowed et al., 2007). Ecological responses to these fluctuations depend on their amplitude and duration compared to the sensitivity and length of species life cycles. Exact ecological responses to physical perturbations will also depend on their temporal position in longer period events and species adaptive options. The slower and more persistent ecosystem changes indexed by OM1 and TVC1 present a large, but poorly defined, range of ecosystem adaptive options, including all those adaptive options available when OM2 and TVC2 change. Changes in OM2 and TVC2 will indicate more rapid changes in the environment characterized by various environmental processes, such as sea temperature change, changes in ocean current speed and direction and changes in productivity. The response of TVC2 will

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depend on the life cycles and migration patterns of the species that have relatively high loading in EOF2 (Fig. 2A), as well as many other ecosystem components that vary similarly. Longer TVC2 periods may indicate local or regional adaptation in temporarily resident populations (Norton, 1999). Smaller species that become sexually mature in their first year, such as anchovies, sardine and market squid, might adapt physically to the changing environment, but for TVC2, migration is a larger factor, particularly for longer-lived species, such as tunas, swordfish and white seabass. The southern California purse seine fishery failure of the early 1920s, discussed in Section 4.2, is most easily interpreted as a TVC2 variation. This relatively short ecosystem fluctuation was characterized by large changes in commercial species availability, but the rebound shows that it did not bring a large change in ecosystem climate; it was an ocean perturbation similar to ones shown by cycling variables in Fig. 2B. Sampling frequency and series length limit this study’s results and uses. Event lengths of more than 40 years for EOF1 and OM1 and less than 20 years for EOF2 and OM2 will probably be conspicuous coupled physical environmental and ecological periods when annual averages are examined in a 50e200 year context. The results indicate regularity in this context, but the unrelated (orthogonal) relationship of EOF1 and EOF2 (Fig. 2), may make their combined presence appear random or chaotic, particularly in the sparsely measured environments. Cyclic patterns in ecological and physical modes indicate forcing by the physical environment or indicate other variables that force nearly simultaneous changes in both of them. However, the largescale and therefore inclusive nature of the physical modes developed here are likely to include most co-occurring processes. Cycles longer than those represented by TVC1 and OM1 or apparent trends, are frequently encountered in environmental data sets (Mantua et al., 1997; Yuan et al., 1997; Finney et al., 2002; Field et al., 2006). Our methods can adjust physical variables for apparent secular trends in ecosystem response (Section 3.4). The current study puts California ecosystem variations into contexts that include marginal fisheries from 1850 to 1905, industrial fisheries from 1920 to 1950, partial moratorium on sardine harvests from 1967 to 1985 and regulated fisheries from 1986 to 2004 (Fig. 5). Current regulations affect the harvest of most of the indicator species (Table 1). However, Fig. 5 suggests a strong physical environmental component in ecosystem (TVC1-proxy) and sardine variability through the 1877e2004 period. Both variables in Fig. 5 have maxima around 1900, but the TVC1-proxy reaches the maximum before OM1. This mismatch of maxima may present a case of complex association where the physical variable continues to increase after the biological variable reached a maximum value a few years before, but it is also possible that the mismatch is caused by variable temporal inconsistencies in the independently derived series. The values for OM1 become progressively more accurate as the global set of SST measurements, from which it was derived, increase in numbers of observations from 1856 to 1981(Kaplan et al., 1998). In addition, the SDR biomass estimates have estimated chronological errors of positive or negative three years (Schimmelmann et al., 1990; Field et al., 2009). Some short-term divergences between the two curves shown in Fig. 5 may represent changes in the spatial distribution of the sardine population that add variability to the early part of the TVC1-proxy record (Field et al., 2009). After 1985, when the biomass estimates are most accurate (Hill et al., 2006), OM1 variations consistently lead TVC1-proxy variations, suggesting causality. Series of sardine abundance estimates from Santa Barbara Basin cores extend over more than 1000 years (Soutar and Isaacs, 1974; Baumgartner et al., 1992; Field et al., 2009). Temporal resolution

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over 1000-year periods is improving (Schimmelmann et al., 2006) and it may become possible to examine interannual and decadal events through these intervals. However, more can be learned about the historical periods by examining the interval between 1850 and 1910. There is correspondence in the variability of OM1 index and TVC1-proxy before 1877 (Fig. 5) and because of agreement in the variables after 1877 it may be possible to interpret the early maximum in sardine biomass. During the sardine maximum in the mid-1800s (Fig. 5), immigrant fishers were drawn to California from China, Japan and European countries. They found a seemingly inexhaustible supply of sardines that were dried and salted for domestic use and export (Holder, 1915; Lydon, 1985; Love, 2006), but these preservation techniques were not sufficient to exploit the large biomass of sardines that was intermittently present along the California coast. The great west coast sardine fishery of the 1930s and 1940s is associated with the second of two early high abundance events (Fig. 5). The dip in sardine abundance between the first two maxima and the associated changes in the ecosystem lasting 10e20 years, were seen as species on the right side of the ecospace became less abundant in the late 19th century. Cries of over fishing and reform were published (Holder, 1915). These warnings appear related to this period of ecosystem change between the two sardine maxima near the end of the 19th and beginning of the 20th centuries. However, it is unlikely that boats, nets and investment in infrastructure of the day were sufficient to change the abundance of fish, except occasionally within areas near major ports. 5.3. California commercial fisheries and environmental variability Commercial fishing businesses would prefer that the weight of valuable fish landed increase, on average, from one year to the next. However, Fig. 2A shows that this has not happened in California. The landings weight did increase from 1930 to 1940, but decreased thereafter for the following 60 years to about a third of the maximum. Total landings value (fishers’ revenue) was more variable. The total revenue increased from 1930 through 1950, then decreased to a minimum in the next 15 years. Additional maxima in dockside value occurred in 1980 and 1990. These changes in total landing weight and revenue occur because many aspects of the fishery are controlled by the ability of the exploited ecosystem to produce the fish and invertebrate harvested. Markets may have modulated the effects of the environment during the early periods with regulations being a more important modulating factor after the early 1980s value maxima (Fig. 2A). Investment by fishers in the early southern California purse seine fishery reached a peak in 1920 with 100 boats worth approximately 20 $M (Skogsberg, 1925). The fisher’s economic problems came in the three following years, before the rebound of the bluefin tuna and other species after 1925 (Figs. 3B and 4). The target species were scarce and in each successive season the fishers mortgaged a greater percentage of their boats and gear to canneries and banks to continue fishing (Skogsberg, 1925). The cannery manager’s willingness to lend money on the basis of an anticipated catch indicates that markets were holding steady or increasing for the species sought and that the purse seine fishers’ economic difficulties arose from unanticipated shortages of harvestable fish. The fish canneries represented major investments in the fishing industry largely in response to an abundant and readily available sardine resource. In the early 1900s sardines were harvested within a few kilometers of the original canneries. In Monterey, much of the sardine harvest before 1920 was within sight of the canneries (Lindner, 1930; Clark, 1939). The Monterey fishers knew in the early 1920s that there were sardines beyond the preferred fishing grounds near port. If they believed that these were un-harvested

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schools, separate from the ones they were harvesting, then the belief that the resource was inexhaustible is understandable. However, the present study suggests that the population reached maximum size in the early 1900s and the capacity of the ecosystem to support large populations of sardines was declining through the period of greatest landings in the 1930s and early 1940s. Fishers and biologists suspected population declines because boats were traveling farther from port to meet cannery demands. Sardine landings increased in the early 1930s as catch per unit of effort decreased (Lindner, 1930; Clark, 1939; Marr, 1960). By 1939 the entire California coast south of Point Arena was part of the harvest area (Clark, 1939). The persistence and growth of cannery and fishery investments through the 1930s and 1940s was due to continued presence of a profitable resource that was harvested more efficiently with improved fishing technology (Herrick et al., 2006). Value came from canning for human food and dehydrating and grinding for high protein fish meal and oil. This balance of costs of harvest versus value held until the late 1940s when the sardine industry was in decline (Marr, 1960; Murphy, 1966), following the decline in the sardine favorable environment and ecosystem (Figs. 3 and 5). California Department of Fish and Game biologists warned the industry, State legislators, and fishery regulators of the decreasing ability of the sardine resource to sustain itself. Some biologists thought the decline was due to changes in the ocean environment (Clark, 1939; Marr, 1960; Herrick et al., 2006). Because the biologists’ warnings were largely ignored, over fishing is also blamed for loss of the resource, but it is not entirely clear that the resource would have been conserved and the fishery sustained by increased regulation. The OM1 curve continued to slope down from the 1900s to the early 1970s suggesting a continued loss of the sardine favorable ecosystem off California (Fig. 5). This continued degradation of sardine favorable environment is similar to the environmental changes of the late 19th century that caused rapid reduction of the sardine populations in the absence of an industrial fishery (Fig. 5). 6. Conclusions Local and large-scale physical to ecological to economic linkages appear similar throughout the 128-year study period. There appear to be definable and quantifiable linkages that operate through wellseparated perturbation cycles of less than 20 years and more than 40 years. There have been several iterations of the less than 20-year perturbations. However, this report has discussed two multidecadal perturbation cycles exceeding 40 years. In this case detailed generalization will require additional study. Empirical orthogonal function analysis of 85-year series of commercial fish and invertebrate landings records leads to the ecospace ecological climate model, which shows variation in species landings maxima through time. When there are changes in the proportion of dominant species in California landings, the position in ecospace changes which indicates changes in the fishers’ harvest opportunities and changes in the ecosystem providing the harvest. Changes in the physical environment corresponding to changes in ecospace, suggest a definite, but presently poorly defined, sequence of events linking the physical environment to the fishers’ economic opportunities. In addition we provide the following notes and conclusions. (1) The first ocean mode (OM1) appears to be an adequate index for large-scale processes that affect the California Current and its ecosystems on multi-decadal cycles longer than 40 years. This mode of environmental variability, which is common over much of the Pacific basin, affects the California Current

(2)

(3)

(4)

(5)

ecosystem and the fisheries that depend on it through local physical and biological processes. Fish and invertebrate species with varying trophic associations harvested from different habitats may have similar, opposite or more complex inter-species population adjustments to the physical and biological environments of their ecosystem. The orthogonal relationship of population adjustments by an array of species is shown in this study. Investment in California fisheries have been initiated and augmented during periods of anticipated high abundance. Economic downturns appear related to environmental changes affecting the ecosystem’s composition. Intensive fishery investment may exacerbate economic downturns. The California Department of Fish and Game commercial fish and invertebrate data, available through the CACom data base, is the only presently known data set of samples directly from the California ocean ecosystem that will support a multispecies investigation that spans 85 years. Marine ecosystem management must accommodate changes in strategy as ecosystems and associated economic and social systems change. These studies indicate some of the ecosystem and economic changes that can be anticipated.

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