Primary Production by Phytoplankton and Microphytobenthos in Estuaries

Ecologists and biogeographers routinely categorize species as "native" with a confidence that for a surprising number of taxa belies the quality of the supporting data. In classical views of the origins of community assemblages, species are either native (that is, indigenous or endemic taxa, including prehistorical invasions) or exotic (that is, historical invasions, including both natural range expansions and human-mediated introductions). The default in this view is to classify species without any obvious record of introduction as "native." Given, however, the historical prevalence of the global movement of plants and animals by humans, combined with a lack of data that would either support or dismiss such movements for many individual species, it is clear that in communities that have been under the sustained influence of human endeavor there are many species that cannot reliably be assigned to either category. The members of this latter group are called "cryptogenic species," a term I introduced in 1982 in abstract form and that has been occasionally but informally used in the literature since that time (e.g., Kinzie 1984). I here define a cryptogenic species (crypt-, Greek, kryptos, secret; -genic, New Latin, genic, origin) as a species that is not demonstrably native or introduced. The word "cryptogenic" (= cryptogenetic) also appears rarely in older medical literature, and was originally, curiously enough, "applied to those cases in which the port of entry of the germs is no longer visible (for example, a hypodermic puncture), or cannot be positively determined" (Oxford English Dictionary 1972). In this note I make two points: (1) cryptogenic species may be remarkably common, and (2) the existence of cryptogenic species has important consequences for understanding biological invasions.

Biological Invasions and Cryptogenic Species

Phytoplankton blooms are prominent features of biological variability in shallow coastal ecosystems such as estuaries, lagoons, bays, and tidal rivers. Long-term observation and research in San Francisco Bay illustrates some patterns of phytoplankton spatial and temporal variability and the underlying mechanisms of this variability. Blooms are events of rapid production and accumulation of phytoplankton biomass that are usually responses to changing physical forcings originating in the coastal ocean (e.g., tides), the atmosphere (wind), or on the land surface (precipitation and river runoff). These physical forcings have different timescales of variability, so algal blooms can be short-term episodic events, recurrent seasonal phenomena, or rare events associated with exceptional climatic or hydrologic conditions. The biogeochemical role of phytoplankton primary production is to transform and incorporate reactive inorganic elements into organic forms, and these transformations are rapid and lead to measurable geochemical change during blooms. Examples include the depletion of inorganic nutrients (N, P, Si), supersaturation of oxygen and removal of carbon dioxide, shifts in the isotopic composition of reactive elements (C, N), production of climatically active trace gases (methyl bromide, dimethylsulfide), changes in the chemical form and toxicity of trace metals (As, Cd, Ni, Zn), changes in the biochemical composition and reactivity of the suspended particulate matter, and synthesis of organic matter required for the reproduction and growth of heterotrophs, including bacteria, zooplankton, and benthic consumer animals. Some classes of phytoplankton play special roles in the cycling of elements or synthesis of specific organic molecules, but we have only rudimentary understanding of the forces that select for and promote blooms of these species. Mounting evidence suggests that the natural cycles of bloom variability are being altered on a global scale by human activities including the input of toxic contaminants and nutrients, manipulation of river flows, and translocation of species. This hypothesis will be a key component of our effort to understand global change at the land-sea interface. Pursuit of this hypothesis will require creative approaches for distinguishing natural and anthropogenic sources of phytoplankton population variability, as well as recognition that the modes of human disturbance of coastal bloom cycles operate interactively and cannot be studied as isolated processes.

Phytoplankton bloom dynamics in coastal ecosystems: A review with some general lessons from sustained investigation of San Francisco Bay, California

In many coastal plain estuaries light attenuation by suspended sediments confines the photic zone to a small fraction of the water column, such that light limitation is a major control on phytoplankon production and turnover rate. For a variety of estuarine systems (e.g. San Francisco Bay, Puget Sound, Delaware Bay, Hudson River plume), photic-zone productivity can be estimated as a function of phytoplankton biomass times mean irradiance of the photic zone. Net water column productivity also varies with light availability, and in San Francisco Bay net productivity is zero (estimated respiratory loss of phytoplankton balances photosynthesis) when the ratio of photic depth ( Z p ) to mixed depth ( Z m ) is less than about 0.2. Thus whenever Z p : Z m Much of the spatial and temporal variability of phytoplankton biomass or productivity in estuaries is explained by variations in the ratio of photic depth to mixed depth. For example, phytoplankton blooms often coincide with stratification events that reduce the depth of the surface mixed layer (increase Z p : Z m ). Shallow estuarine embayments (high Z p : Z m ) are often characterized by high phytoplankton biomass relative to adjacent channels (low Z p : Z m ). Many estuaries have longitudinal gradients in productivity that mirror the distribution of suspended sediments: productivity is low near the riverine source of sediments (low Z p : Z m ) and increases toward the estuary mouth where turbidity decreases. Some of these generalizations are qualitative in nature, and detailed understanding of the interaction between turbidity and estuarine phytoplankton dynamics requires improved understanding of vertical mixing rates and phytoplankton respiration.

Turbidity as a control on phytoplankton biomass and productivity in estuaries

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.

https://science.sciencemag.org/content/sci/231/4738/567.full.pdf

The Modification of an Estuary

Phytoplankton dynamics in the upper reach of the northern San Francisco Bay estuary are usually characterized by low biomass dominated by microflagellates or freshwater diatoms in winter, and high biomass dominated by neritic diatoms in summer. During two successive years of very low river discharge (the drought of 1976-77), the summer diatom bloom was absent. This is consistent with the hypothesis that formation of the diatom population maximum is a consequence of the same physical mechanisms that create local maxima of suspended sediments in partially-mixed estuaries: density-selective retention of particles within an estuarine circulation cell. Because the estuary is turbid, calculated phytoplankton growth rates are small in the central deep channel but are relatively large in lateral shallow embayments where light limination is less severe. When river discharge falls within a critical range (100–350 m3 s−1) that positions the suspended particulate maximum adjacent to the productive shallow bays, the population of neritic diatoms increases. However, during periods of high discharge (winter) or during periods of very low discharge (drought), the suspended particulate maximum is less well-defined and is uncoupled (positioned downstream or upstream) from the shallow bays of the upper estuary, and the population of neritic diatoms declines. Hence, the biomass and community composition of phytoplankton in this estuary are controlled by river discharge.

River discharge controls phytoplankton dynamics in the northern San Francisco Bay estuary

Detailed surveys throughout San Francisco Bay over an annual cycle (1980) show that seasonal variations of phytoplankton biomass, community composition, and productivity can differ markedly among estuarine habitat types. For example, in the river-dominated northern reach (Suisun Bay) phytoplankton seasonality is characterized by a prolonged summer bloom of netplanktonic diatoms that results from the accumulation of suspended particulates at the convergence of nontidal currents (i.e. where residence time is long). Here turbidity is persistently high such that phytoplankton growth and productivity are severely limited by light availability, the phytoplankton population turns over slowly, and biological processes appear to be less important mechanisms of temporal change than physical processes associated with freshwater inflow and turbulent mixing. The South Bay, in contrast, is a lagoon-type estuary less directly coupled to the influence of river discharge. Residence time is long (months) in this estuary, turbidity is lower and estimated rates of population growth are high (up to 1–2 doublings d−1), but the rapid production of phytoplankton biomass is presumably balanced by grazing losses to benthic herbivores. Exceptions occur for brief intervals (days to weeks) during spring when the water column stratifies so that algae retained in the surface layer are uncoupled from benthic grazing, and phytoplankton blooms develop. The degree of stratification varies over the neap-spring tidal cycle, so the South Bay represents an estuary where (1) biological processes (growth, grazing) and a physical process (vertical mixing) interact to cause temporal variability of phytoplankton biomass, and (2) temporal variability is highly dynamic because of the short-term variability of tides. Other mechanisms of temporal variability in estuarine phytoplankton include: zooplankton grazing, exchanges of microalgae between the sediment and water column, and horizontal dispersion which transports phytoplankton from regions of high productivity (shallows) to regions of low productivity (deep channels).

Temporal dynamics of estuarine phytoplankton: A case study of San Francisco Bay

............................................................... 6 Introduction............................................................ 7 Methods................................................................. 8 References.............................................................. 11

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Raymond L. J. Wong

Papers

River discharge controls phytoplankton dynamics in the northern San Francisco Bay estuary

Temporal dynamics of estuarine phytoplankton: A case study of San Francisco Bay

Plankton studies in San Francisco Bay, California; XIV, Chlorophyll distributions and hydrographic properties of San Francisco Bay, 1992

Plankton studies in San Francisco Bay; II, Phytoplankton abundance and species composition, July 1977-December 1979