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15 ’ , He Continued, 1890 1910 1930 1950 1970 1990 YEAR MACCALL: LOW-FREQUENCY VARIABILITY IN FISH POPULATIONS CalCOFl Rep., Vol. 37, 1996 PAllERNS OF LOW-FREQUENCY VARIABILITY IN FISH POPULATIONS OF THE CALIFORNIA CURRENT ALEC I). MACCALL N~tion~lManne Fi5herim Service South\\ect Fi5heries Science Chter 31 50 Paradise Ilr. Tihuroii, C::ilifwtiia 94c120 ABSTRACT kinds of conditions. Rather, there are probably a great Long-term data sets include sedimentary fish-scale number of possible regimes and abrupt discontinu- paleochronologies, faunal surveys froiii the 1SOOs, and ities connecting them, flip-flops froin one regime to the records of game fish catches maintained by the Tuna another; niultifarious regimes involving biology or Club at Avalon since 1898. These data suggest a inode climate, or oceanography, or migrations, temperature, of low-frequency biological variability over a cycle of or weather, or combinations of these . My main .5O to 70 years that is associated with alternation of warm point is that there are no simple statistics in the ordi- and cold physical regimes. The very warm conditions nary sense. There are internal, interactive episodes in the northeastern Pacific Ocean since 1976 seein sim- locked into persistence, and one is entirely fooled if ilar to conditions experienced from about 1850 to 1870. one takes one of these short intervals of a decade or Coastal pelagic fish abundances since 1930 indicate an so and decides there is some sort of simple probabil- orderly sequence of four or five dominant species through ity associated with it. organisms must respond to these cycles that is very similar to the order and timing more than just fluctuations around some optimum of fluctuations in catches of siniilar pelagic fishes in condition. Actually, many of their characteristics and Japanese waters. The consistency arid predictability of fluctuations of populations must be related to these this rotation and the biological relationship to low- very large alternations of conditions. frequency fluctuations in physical conditions are iin- Isaacs coined the term wyim to describe distinct en- portant areas for further research. These patterns of low- vironmental or climatic states, and yqime SI$ has recently frequency variability result in boom-and-bust fisheries, gained acceptaiice as a term for the abrupt transitions and pose serious probleiiis for “sustainable development.” between regimes. It is notable that publication of lsaacs’ regime concept in 1976 itself coincided with a profound INTRODUCTION reginie shift in the northeastern Pacific. That shift was With recognition of the worldwide decline in fish not apparent at the time, and it required another decade stocks, the guideline of “sustainable development” is re- for the oceanic warming (figure 1) to be recognized as ceiving international support as an explicit goal of fish- a new and persistent condition (MacCall and Prager ery and ecosystem management. Ironically, this conies 1 988; Trenberth 1990). at a time when the biological basis of sustainability in In ths paper I exanine variability in southern Cahfornia some of the world’s niajor fisheries is itself being ques- tioned because of natural patterns of variability (e.g., Lluch-Belda et al. 1989; Beaniish 1995). In a broader 19 , context, the science of ecology is moving away froni Fta- tic and equilibrium views of ecosystems. Rather, scien- tists are coining to perceive ecosystems as pulsing and fluctuating entities, driven by external sources of vari- ability and by internal nonlinear dynamics. Odurii et al. (1995) state the possibility that “what is sustainable in ecosystems, is a repeating oscillation that is often poised on the edge of chaos.” At the 1973 CalCOFI Symposium, John Isaacs (1976) took issue with the common implicit assumption in fish- eries science that there is “some steady state . around which there is a normal distribution of perturbations.” 15 ’ , He continued, 1890 1910 1930 1950 1970 1990 YEAR This certainly does not seein to be the case. The as- Figure 1. Mean annual temperatures (“C) at the Scripps Institution of sumption is that there are some normal statistics to all Oceanography Pier, La Jolla, California. 1 00 MACCALL: LOW-FREQUENCY VARIABILITY IN FISH POPULATIONS CalCOFl Rep., Vol. 37, 1996 fish populations over the longest time scales supported The major warming that began about 1976 is clearly by the qualitative and quantitative information available. visible in the Scripps Pier temperature record. This event Even the quantitative time series are niuch too short for was widespread and occurred over the entire northeast- rigorous statistical analysis. My approach to these data ern Pacific Ocean, from Mexico to Alaska (Cole and is that of “historical science” (Gould 2989; Francis and McLain 1 989). However, the geographic extent of ear- Hare 1994); consequently, some of my interpretations lier temperature regimes and regime shifts is less clear. are speculative. An earlier regime shift occurred about 1940, begin- Perhaps because fish are an important commodity, ning a long period of cold temperatures. Consequently better long-term records exist for fishes than for any the Scripps Pier record consists of three conspicuous other biological component of the California Current temperature regimes: a moderate and highly variable pe- System. Paleochronologies of fish debris preserved in riod fiom the beginning of the time series to about 1940, anaerobic sediments provide invaluable information on a cold period from 1940 to 1976, and a warm period prehistoric fluctuations and very-low-frequency vari- since 1976. Ware (1995) describes similar alternations of ability. The earliest historical records indicative of fish temperature regimes off British Columbia. abundance in the California Current are relatively qual- It is not clear how these low-frequency temperature itative and begin with the Pacific Railroad Survey of regimes are related to the El Nifio/Southern Oscillation. 1856. Quantitative big game fishing records have been However, brief warm events related to both El Niiio and maintained since the turn of the century by the Tuna variability in the Aleutian Low (Wooster and Hollowed Club at Avalon, on Santa Catalina Island. Although Tuna 1995) are superimposed on these underlying tenipera- Club data have seen little use in scientific contexts, these ture regimes. Major warm events have occurred at a tran- records constitute the longest quantitative and unbroken sition (1940), in the midst of a cool period (1958-.59), series of biological observations in this region. The State and in the midst of a warm period (1982-83 and of California began keeping systematic records of fish 1992-93). landings in 191 6, and began comprehensive biological sampling of coastal pelagic fish landings in the 1930s. FISH ABUNDANCES Good information on ichthyoplankton exists for niuch The anaerobic sediments of the Santa Barbara Basin of the last 50 years due to the California Cooperative provide a remarkable paleochronology of fish scale-de- Oceanic Fisheries Investigations (CalCOFI) and other position rates (Soutar 1967). Soutar and Isaacs (1974) fishery and oceanographic programs that were initiated developed a 160-year time series of Pacific sardine following World War 11. (Savdirzopr sqax) and northern anchovy (Engraulis ~OY- For the purpose of this paper, I use ocean tempera- dux) scale-deposition rates that indicated large natural ture as a simple proxy for the more complicated and long-term fluctuations in both of those species. poorly understood suite of oceanographic variables that Baumgartiier et al. (1‘992) extended and refined these influences biological populations. Sea-surface tenipera- studies, and developed a paleochronology spanning the tures have been monitored continuously at the Scripps last 1,700 years at a temporal resolution of 30 years Institution of Oceanography Pier (Scripps Pier) in La (figure 2). Power spectra of these long time series show Jolla, beginning in 1916 (figure I), forming one of the high variability in both sardine and anchovy scale-de- longest time series of sea-surface temperatures on the position rates at a period of approximately GO years, Pacific coast. Earlier observations of sea-surfxe tem- and at longer periods that differ for the two species. perature from ships-of-opportunity are provided by the Ware’s (1995) analysis of a much shorter physical and bi- COADS data set (Mendelssohn and Roy l996), but there ological data set also indicated a spectral peak in the range are insufficient observations from southern California of 50 to 75 years. Neither Baunigartner et al. (1992) nor waters before 1920 to provide annual resolution of teni- Ware (1995) was able to identify a physical mechanism perature fluctuations (K. Parrish, PFEG, NMFS, Pacific associated with this low-frequency variability. Grove, CA, pers. conini. 9/13/95). However, Parrish’s examination of the post-1920 COADS data did denion- strate that low-frequency temperature fluctuations in Anecdotal Information southern California do not necessarily parallel temper- Biological surveys of the Pacific coast began in the ature fluctuations off central or northern California. niid-l80Os, and provide useful indications of presence Thus, annual average Scripps Pier temperatures may pro- or absence of species but have poor resolution for the vide a siniplified representation of southern California purpose of estimating the timing and duration of ,envi- ocean conditions but may not always be a reliable indi- ronmental regimes. However, in comparison with con- cator of conditions in adjacent Pacific coastal regions ditions during most of the twentieth century, some of such as central California. the early survey information is so peculiar that Hubbs 101 MACCALL LOW-FREQUENCY VARIABILITY IN FISH POPULATIONS CalCOFl Rep., Vol. 37, 1996 PACIFIC SARDINE NORTHERN ANCHOVY 56 -- J(1 -- T 15 20 15 10 5 0 xx) xg a Ma 700 800 900 loo0 1100 1200 1JM) 1400 1Mo 16W 17W 1800 1803 2000 YEAR Figure 2. Sardine and anchovy scale-deposition rates in the Santa Barbara Basin (reproduced from Baumgartner et al. 1992) (1948) was able to infer the existence of an unusually California until a small population reappeared in San warm ocean climate during the mid 1800s.
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