The San Francisco Bay estuary has been rapidly modified by human activity. Diking and filling of most of its wetlands have eliminated habitats for fish and waterfowl; the introduction of exotic species has transformed the composition of its aquatic communities; reduction of freshwater inflow by more than half has changed the dynamics of its plant and animal communities; and wastes have contaminated its sediments and organisms. Continued disposal of toxic wastes, the probable further reduction in freshwater inflow, and the possible synergy between the two provide the potential for further alteration of the estuary's water quality and biotic communities.
Long-term macrobenthic sampling at a site in northern San Francisco Bay has provided an unusual opportunity for documenting the time course of an invasion by a recently introduced Asian clam Potamocorbula arnurensis. Between 1977, when sampling began, and 1986, when the new clam was first discovered, the benthic commun~ty varied predictably in response to river inflow. During years of normal or high river inflow, the community consisted of a few brackish or freshwater species. During prolonged periods of low river inflow, the number of species doubled as estuarine species (e.g. Mya arenana) migrated up the estuary. In June 1987, at the beginning of the longest dry period in recent decades, large numbers (> 12 000 m-') of juvenile P. arnurensis were discovered at the site. By midsummer 1988 the new clam predom~nated (> 95 %) in both total number of individuals and biomass, and the expected dry-penod estuarine species did not become re-established. The rapid rise of P. arnurensls to numerical dominance throughout the region of the original introduction was probably facilitated by the fact that this region of the bay had been rendered nearly depauperate by a major flood in early 1986. Once introduced, the clam had sufficient time (> 1 yr) to become well established before the salinity regime was appropriate for the return of the estuarine species. Subsequently, the new clam was apparently able to prevent the return of the dry-period community. Its ability to live in low salinity water (< 1 %o) suggests that P. amurensis may not be displaced with the return of normal winter river flow and, therefore, may have permanently changed benthic community dynamics in this region of San Francisco Bay.
The benthic invertebrate community inhabiting the extensive and sedimentologically homogeneous mudflats of South San Francisco Bay has demonstrated a high degree of constancy in both species composition and relative abundance among species throughout 10 yr of observation. The community, composed predominantly of introduced species with opportunistic lifestyles, is dominated numerically by Gemma gernma, Ampelisca abdita, and Streblospio benedicti. The key to the persistent CO-occurrence of these species on the mudflats seems to lie in the combination of (1) the recurrence of minor disturbances of the mudflat habitat (e.g. sediment depositlon/erosion, inundation by low-salinity water) on time scales comparable to that of life cycles; (2) opportunistic life history strategies (rapid maturity, brooding of young, multiple generations each year, ease of local dispersal of both juveniles and adults) that permit continued colonization of the mudflat surface or rapid recolonization after disturbances. Only 1 of the 3 numerically-dominant species. A. abdita, displays an annual periodicity in abundance. S. benedicti and G. gemma, through broadly flexible reproductive strategies permitted in the mild San Francisco climate, can exhibit strong recruitment at any time between spring and autumn. The most extreme community changes, involving temporary reduction or elimination of normally dominant populations, occurred as a result of anomalous disturbances such as unusual buildup and decay of an algal mat during 1 summer and prolonged periods of unusually high freshwater inflow during 2 successive winters. The introduced opportunists routinely CO-occur at high densities. However, one of these, the tube-dwelling amphipod A. abdita, may control the abundance of the native mollusk Macoma balthica.
The von Bertalanffy model of body growth is inappropriate for organisms whose growth is restricted to a seasonal period because it assumes that growth rate is invariant with time. Incorporation of a time-varying coefficient significantly improves the capability of the von Bertalanffy equation to describe changing body size of both the bivalve mollusc Macoma balthica in San Francisco Bay and the flathead sole, Hippoglossoides elassodon, in Washington state. This simple modification of the von Bertalanffy model should offer improved predictions of body growth for a variety of other aquatic animals. Key words: Bertalanffy, growth model, growth rate, Macoma balthica
Results from multi-year investigations in the San Francisco Bay estuary show that large abundance fluctuations within benthic macroinvertebrate populations reflect both (1) within-year periodicity of reproduction, recruitment, and mortality that is not necessarily coincident with seasonal changes of the environment (e.g., the annual temperature cycle), and (2) aperiodic density changes (often larger than within-year fluctuations) following random perturbations of the environment.Density peaks of the small, short-lived estuarine invertebrates that comprise the vast majority of individuals in the bay's relatively homogeneous benthic community normally occur between spring and autumn depending on the species, in large part a reflection of reproductive periodicity. However, because mild winters permit reproductive activity in some of the common species throughout much of the year, other factors are important to within-year density fluctuations in the community. Seasonally predictable changes in freshwater inflow, wind and tidal mixing, microalgal biomass, and sediment erosion/deposition patterns all contribute to observed seasonal changes in abundance. For example, the commonly observed decline in abundance during winter reflects both short-lived species that die after reproducing and the stress of winter conditions (e.g., inundation by less saline, sediment-laden water and the decline in both planktonic and benthic algal biomass -a direct source of food for the shallow-water benthos). On the other hand, data from several studies suggest that observed 'recruitment' and 'mortality' may in fact be the migration of juveniles and adults to and from study sites. For example, the common amphipod Ampelisca abdita apparently moves from shallow to deep water, or from up-estuary to down-estuary locations, coincident with periods of high river runoff in winter. Growth of individuals within the few studied species populations is also highly seasonal, and appears to be coincident with seasonal increases in the abundance of planktonic and/ or benthic microalgae.Two multi-year studies have shown that, in addition to within-year periodicity, major restructuring of the benthic community can occur as a result of anomalous (usually climate-related) perturbations of the benthic habitat. For example, during wet years freshwater-intolerant species disappear from the upper part of the estuary and from shallow areas of the bay. During a two-year drought these same species colonized the extreme upper end of the estuary in large numbers. Other aperiodic perturbations include localized instances of sediment erosion or deposition and algal mat accumulations that greatly depress abundance. Additionally, there is evidence (observations that the clam Macoma balthica establishes large populations only when the amphipod A. abdita is not abundant) that species interactions can contribute greatly to interannual variations. Thus, while community composition may change little over the long term, year-to-year predictability of species abundances is low.
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