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THE DN- Columbia River Estuary

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THE DN- Columbia River Estuary - - ;; . So no is e ~~~~~FOR DISPLAY ONLY THE DN- Columbia River Estuary Data Development Program (CREDDP) THE DYNAMICS OF THE COLUMBIA RIVER ESTUARINE ECOSYSTEM VOLUME II June 1984 THE DYNAMICS OF THE COLUMBIA RIVER ESTUARINE ECOSYSTEM AUTHORS Charles Simenstad David Jay C. David McIntire Willa Nehlsen Christopher Sherwood Lawrence Small GRAPHICS COORDINATION WORD PROCESSING Krebill Isabel Turner Elizabeth William Barnett Julie Guerrero Elizabeth Rummell TABLE OF CONTENTS (VOLUME II) Page INTEGRATION 341 6. CONCEPTUAL FRAMEWORK FOR PHYSICAL-BIOLOGICAL 341 6.1 THE PROBLEM OF INTEGRATION 343 6.2 THE PROCESS CONCEPT 345 6.3 AN ECOSYSTEM MODEL OF ESTUARINE PROCESSES 350 Primary Food Processes 6.3.1 351 6.3.2 Subsystems of Primary Food Processes 352 6.3.3 Consumption 353 6.3.4 Subsystems of Consumption 354 6.4 SUMMARY 363 7. ECOSYSTEM PROCESSES 363 7.1 PRIMARY FOOD PROCESSES: PRIMARY PRODUCTION 363 Introduction 7.1.1 371 7.1.2 Taxonomic Structure of Plant Assemblages 384 7.1.3 Biomass 7.1.4 Resource Inputs and Controlling Physical 394 Variables 404 7.1.5 Primary Productivity 428 7.1.6 Process Generated Outputs 433 7.1.7 Non-Process Generated Inputs and Outputs 444 7.1.8 Annual Production Dynamics 450 7.2 PRIMARY FOOD PROCESSES: DETRITAL DECOMPOSITION 450 7.2.1 Introduction 452 7.2.2 Detrital Biomass 7.2.3 Resource Inputs and Controlling Physical Processes 457 458 7.2.4 Process-Generated Outputs Outputs 459 7.2.5 Non-Process Generated Inputs and 463 7.3 CONSUMPTION PROCESSES 463 7.3.1 Introduction 7.3.2 Taxonomic Structure and Organization of 465 Consumer Assemblages 515 7.3.3 State Variables 555 7.3.4 Process Dynamics 590 7.4 SUMMARY 590 7.4.1 Primary Production V Page 7.4.2 Detritus Production 593 7.4.3 Consumption 593 8. ECOSYSTEM ANALYSES BY REGIONS AND HABITAT TYPES 599 8.1 ESTUARINE REGIONS 599 8.1.1 Entrance (Region 1) 600 8.1.2 Baker Bay and Trestle Bay (Region 2) 600 8.1.3 Estuarine Channels (Region 3) 601 8.1.4 Youngs Bay (Region 4) 602 8.1.5 Mid-Estuary Shoals (Region 5) 602 8.1.6 Grays Bay (Region 6) 602 8.1.7 Cathlamet Bay (Region 7) 603 8.1.8 Fluvial Region (Region 8) 603 8.2 ESTUARINE HABITAT TYPES 603 8.3 BIOLOGICAL PRODUCTION BY REGIONS AND HABITAT TYPES 607 8.3.1 Producers 607 8.3.2 Consumers 611 8.3.3 Production Comparisons 616 9. SUMMARY AND CONCLUSIONS 663 9.1 SUMMARY 663 9.1.1 Circulation 663 9.1.2 Sedimentation 664 9.1.3 Primary Production 666 9.1.4 Detritus 668 9.1.5 Primary Consumers 669 9.1.6 Predators 670 9.2 THE RIVER-ESTUARY-PLUME CONTINUUM 670 9.3 CONSEQUENCES OF HISTORICAL CHANGES TO THE ECOSYSTEM 671 9.4 GAPS IN UNDERSTANDING OF ECOSYSTEM AND RECOMMENDATIONS FOR FURTHER RESEARCH 674 9.4.1 Estuarine Circulation and Sedimentation 674 9.4.2 Detritus Sources and Cycling 676 9.4.3 Taxonomic Structure of Phytoplankton and Benthic Microalgal Assemblages 677 9.4.4 Primary Production Processes 677 9.4.5 Import and Export of Fluvial Phytoplankton and Zooplankton 678 9.4.6 Consumption Processes 678 9.5 CONCLUSIONS 678 LITERATURE CITED (VOLUME II) 681 vi 6. CONCEPTUAL FRAMEWORK FOR PHYSICAL-BIOLOGICAL INTEGRATION A major goal of this report is to provide an integrated presentation of Columbia River Estuary community dynamics in relation to physical processes. This chapter presents the conceptual model used in carrying out physical-biological integration. Problems in integrating CREDDP data and considerations in model selection are described (Section 6.1). The modeling approach (Section 6.2) and details of the model itself (Section 6.3) are then presented. The model is discussed according to major biological processes: Primary Food Processes (Primary Production and Detrital Decomposition) and Consumption (Deposit Feeding, Suspension Feeding, Wetland Herbivory, and Predation). 6.1 THE PROBLEM OF INTEGRATION Information available for integration represented many levels of biological organization and of spatial and temporal resolution. For example, the biological data included species lists, counts of organisms and estimates of densities, biomass measurements of species and groups of species, life history information about species, estimates of primary production and respiration, density and flux of organic particles in the water column, and chlorophyll concentrations in the water column and sediment. Furthermore, these data related to distributional patterns and system dynamics within a variety of temporal and spatial frames. The major challenge in integrating CREDDP data was to develop a conceptual model that could accommodate the Program's variety of organizational frameworks and resolution levels. One possible approach was a trophic (food web) analysis of the estuarine community. In this case, estimates of production and energy inputs and outputs for all functional groups of organisms would be required. Unfortunately, such data were lacking for many of the consumer groups. Moreover, data related to complex interactions between detritus and other. biological components of the system also were insufficient for a complete trophic synthesis. For a satisfactory integration of scientific information, it also was necessary to include structural and distributional information in relationship to physical processes. CREDDP produced much data related to temporal and spatial distributions of organisms. Summaries of such data -may be more relevant to certain management problems. than trophic considerations, particularly problems related to the vulnerability of organisms to perturbation at specific locations. Several conceptual models were considered for the synthesis of scientific information from CREDDP. The most notable possibilities included the Fish and Wildlife Service model from the Ecological Characterization of the Pacific Northwest Coastal Region (Proctor et. al. 1980), a trophic level model (Lindeman 1941; 1942), habitat-based model suggested by personnel of the Columbia River Estuary Study Taskforce, a hierarchical process model based on the concepts of Overton (1972, 1975, 1977) and McIntire (1983), and the hypothesis systems 341 PRIMARY CONSUMER PROCESSES EXPORT IMPORT I RESOURCES RESPIRATION PARTICULATE rPHYSICAL \DDISLE PR~~~'CESSES ~~~~~DISSOLVEDAN QPROCESSES 2 < ORGANIC MATTER Figure 6.1. A diagram of a generalized primary consumer process illustrating its state variable and relevant input and output variables. In this case, the process of interest (large circle) is coupled to Predation, a secondary consumer process. 342 Work Unit of CREDDF in models introduced by the Biophysical Interactions February 1980. was similar to that The modeling approach adopted for integration in Section (1975) and McIntire (1983). The model, described of Overton the many resolution 6.2, has a hierarchical structure which accommodates emphasizes biological levels of the CREDDP data. Although the model of important species processes, its structure also permits consideration in these processes. and functional groups of species which are involved organize the presentation Consequently, the model is flexible enough to as information concerning of process dynamics in the estuary as well processes are treated as distributional patterns of species. Physical biological processes. system inputs that limit, control, or regulate 6.2 THE PROCESS CONCEPT physical and ecological literature often refers- to various The intuitively processes in contexts- that are usually biological for without a formal, theoretical structure. However, understandable concept by purposes, it is desirable to formalize the process analytical convention for definitions and the establishment of a explicit apply only to diagramming relationships. Here,- the definitions treated conceptually as biological processes; physical processes are driving or control functions. Definitions: relevant to the (1) A process is a systematic series of actions (or modeled). dynamics of the system as-it is conceptualized represents the net (2) A state variable is a variable that - any instant of accumulations: of a material -of interest, at time. are the (Examples of state variables -in ecological systems species and the biomasses of species or functional groups of concentration of a nutrient.) that potentially (3) A resource is any input required by a process : may be in limited supply. This 6.1 is a generalized diagram of a biological process. Figure to diagram illustrates the six symbols that ,will be used drawing also research processes that were investigated by CREDDP the individual by circles. A projects. Biological processes are always represented of interest, while large. circle is used to focus on a particular process that is coupled to the a smaller circle indicates another -process general structure interest. In the case of Figure 6.1, the process of consumer process the featured process is compatible with a -primary of elaborated into one grazing or deposit feeding). This process is (e.g., outputs and other variables that, represent inputs and state variable- variable, with the process. - In this case,; the -state associated of time that by the rectangle, is the biomass at any instant illustrated that -follow, most is involved in the process. In the system diagrams functional groups of state variables will represent biomasses of various Other variables organisms, associated with a process of interest. include resource inputs, associated with the process in Figure 6.1 343 imports of biomass from outside the spatial area of interest, export of biomass from the system, respiratory and waste losses generated by the process, losses of biomass to the process of predation, and the set of relevant physical variables that influence the -process. In summary, biological processes are defined as those biological activities that are generated by complex interactions among a set of variables, namely the corresponding inputs, outputs and state variables. For the synthesis of biological data from CREDDP, the problem of qualitative differences among organisms involved in a particular process is approached, by partitioning the process biomass into the, number of state variables that corresponds to the taxonomic entities or functional groups of organisms investigated by the various CREDDP projects.
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