2. Food Web Evaluation
This version of the Puget Sound Science Update provides an initial evaluation of food web indicators, but is not intended to be comprehensive. Highlights include the evaluation of individual species or species complexes as food web indicators due to their key functional roles (e.g., forage fish, jellyfish), and the identification of existing data sources for assessing food web structure and function at Washington State agencies and via satellite.
Next Step: Future versions of this document would benefit from the evaluation of more indicators pertinent to the Freshwater and Terrestrial Domains, and the inclusion of more candidate indicators in the Marine Domain to ensure a full treatment of the key attributes identified in Section 3.2.3.3. Indicators of energy and material flows deserve particular attention in future assessments, as they were not the focus of the review by O’Neill et al. (2008). |
Key Point: Because of the difficulty of directly measuring attributes of food web health, ecosystem models have the potential to greatly contribute to the evaluation of foodweb indicators (Samhouri 2009). We encourage the development of ecosystem models as a tool for testing the performance of food web indicators. |
Indicators of community composition
Species abundance and biomass data can be used to paint a synthetic picture of community composition, especially when viewed collectively with respect to particular Domains and in relation to species’ trophic and functional roles. Even in isolation, insight into the status and trends of keystone species (i.e., those that have a disproportionate influence on food web structure relative to their abundance), highly connected species (i.e., those that are consumers of and consumed by many other species), minimally connected species, and those species representing a large proportion of the biomass in Puget Sound can be useful for interpreting the structural configuration of the food web (Soulé et al. 2005). In addition, species abundance and biomass data can be summarized into index values that describe the three different types of diversity defined in Section 3.2.3.3 (species, trophic, and response diversity). Dietary composition data, especially for higher trophic level predators such as marine mammals and birds, offer an alternative inroad to understanding community composition in Puget Sound and are available for a limited subset of species. Ongoing monitoring programs led by the Washington Departments of Ecology, Fish and Wildlife, and Natural Resources, among others, provide a rich source of information on community composition in Puget Sound. The challenge is to sort through these data to extract meaningful summary descriptions.
Indicators of energy and material flows
Proxies for primary productivity such as chlorophyll a concentration and phytoplankton biomass (in the Marine Domain) and leaf area index (in the Terrestrial Domain) are the most widely available indicators for energy and material flows in Puget Sound. Remote-sensing data are a valuable source of this information, though other, labor-intensive approaches are available for obtaining spatially explicit and finely resolved understanding of primary productivity as well (e.g., plankton tows, forest inventories, etc.). Alternatives to remote-sensing data are especially important in the Marine Domain, where it is difficult to obtain reliable estimates of primary productivity in nearshore areas at small spatial scales. More detailed data collection or modeling efforts (e.g., Ecopath with Ecosim) are needed to estimate the magnitude of secondary production and pathways of energy flows throughout the food web. Biogeochemical approaches for measuring cycling rates are well developed, especially with respect to inorganic nutrients, but such data are not widely available and can be quite expensive to obtain. Making up for this deficiency will require detailed, broad-scale studies of how different species interact with the physical and chemical oceanography of Puget Sound to affect processes such as nitrogen fixation, carbon sequestration, and microbial decomposition.
Evaluation of food web indicators in Puget Sound
There were nineteen Food Web indicators identified and of these, we have evaluated fifteen. The degree to which food web indicators satisfy our evaluation criteria is highly variable, and about half of them did not perform well against any of the criteria. The majority of evaluated indicators were from the Marine Domain, and no indicators have yet been evaluated for Freshwater Food Webs. The current status of indicator evaluations for the Food Webs Goal is summarized below.
Marine food web indicator evaluation
Eleven indicators of Marine Food Web community composition and two indicators of Marine Food Web Energy and Material Flows were evaluated (Table 11 and Table 12). The status and trends of benthic and pelagic fish communities species, marine shorebird diets, and jellyfish abundance performed best against the primary considerations for indicators of community composition. Of these indicators, however, only marine shorebird diets also met a majority of the Data and Other Considerations criteria. The general deficiency of quantitative data suggests the potential utility of several indicators while highlighting the need to begin data collection and monitoring. Most of the community composition indicators that did not perform well against the Primary Considerations also were deficient under the Data Considerations criteria. One of the biggest challenges for developing Marine Food Web indicators will be to increase their specificity prior to evaluation; several indicators, like the marine biodiversity index, shellfish, and benthic macroinvertebrates, were considered too vague to evaluate properly.
Phytoplankton biomass and chlorophyll a concentration provide similar information about primary productivity in the Puget Sound Marine Food Web. Both indicators performed well against the Primary Considerations for indicators of energy and material flows. However, chlorophyll a concentration met more of the Data and Other Considerations. Due to this indicator’s reliance on remotely sensed data, however, it is unlikely to provide information about energy and material flows on spatial scales smaller than the PSP Action Areas. We suggest the evaluation of additional indicators of energy and material flows in the future.
Table 11. Summary of Marine Food Webs – Community Composition indicator evaluations. The numerical value that appears under each of the considerations represents the number of evaluation criteria supported by peer-reviewed literature. For example, Macro benthic inverts has peer-reviewed literature supporting 0 out of 5 Primary Considerations criteria. Details can be found in the accompanying spreadsheets.
Guild |
Indicator |
Primary Considerations (5) |
Data Considerations (8) |
Other Considerations (5) |
Summary Comments |
Mammals |
Harbor seals – food web interactions |
1 |
5 |
2 |
Should be a good indicator of fish community composition, and possibly of population condition. Breadth of seal diet may limit power to detect small changes. Spotty historical data available throughout the region. |
Key Fish |
Benthic fish species status & trends |
4 |
3 |
0 |
Overall appear to be good indicators of food web community composition, although historical data is currently lacking making it difficult to determine long-term trends. |
|
Bentho-pelagic fish status & trends |
4 |
2 |
0 |
Overall appear to be good indicators of food web community composition, although historical data is currently lacking making it difficult to determine long-term trends. |
|
Bottomfish species (rats and flats) status & trends |
1 |
0 |
4 |
Bottomfish noted as best indicator for some ecosystem attributes [118], although only appears as adequate indicator using our criteria. Difficult to determine if this indicator responds predictably to ecosystem attribute or actions/pressures. Patchy historical data. |
Birds |
Marine shore birds – food web interaction |
3 |
7 |
2 |
Overall a good indicator, with relevance to forage fish prey species (diet variability responds to prey variability). Historical data available, although limited to two PSP action areas. |
Shellfish & Other Inverts |
Jellyfish |
4 |
3 |
2 |
Theoretically sound – jellyfish should be reliable indicators of trophic energy transfer & community composition. Responds predictably to actions and pressures, and may be especially relevant to understanding the status of forage fish. Historical data is limited, although still a promising indicator. |
|
Shellfish |
0 |
0 |
0 |
Currently unable to properly evaluate because indicator is too vague. Recommend selection of particular species of bivalves as indicators. |
|
Macro benthic inverts |
0 |
0 |
0 |
Currently unable to properly evaluate because indicator is too vague. Recommend selection of particular species of benthic inverts as indicators. |
Key Species |
Marine biodiversity index |
0 |
0 |
0 |
Currently there is not sufficient information available to evaluate this indicator; the WA Biodiversity Council has planned to develop this indicator further. |
|
Marine fish/invert status & trends in marine reserves |
0 |
0 |
0 |
Consolidate this indicator with ‘marine fish/invert status & trends at rocky habitats’. If monitored inside marine reserve, it should also be monitored outside reserve to serve as a reference point. |
|
Marine fish/invert status & trends at rocky habitats |
1 |
4 |
0 |
Difficult to evaluate as currently defined; need to explicitly define species or community parameters of interest. Some historical data available. |
Table 12. Summary of Marine Food Webs – Energy and Material Flow indicator evaluations. The numerical value that appears under each of the considerations represents the number of evaluation criteria supported by peer-reviewed literature. For example, Chlorophyll a has peer-reviewed literature supporting 3 out of 5 Primary Considerations criteria. Details can be found in the accompanying spreadsheets.
Guild |
Indicator |
Primary Considerations (5) |
Data Considerations (8) |
Other Considerations (5) |
Summary Comments |
Plants |
Phytoplankton biomass |
3 |
1 |
1 |
Good indicator of pelagic ecosystems, especially nutrient cycling and the amount of primary production. Only limited amounts of historical data available. Provides similar information as chl a so choose one to avoid redundancy. |
|
Chlorophyll a |
3 |
7 |
2 |
Chl a is a good proxy for overall primary productivity and nutrient cycling/uptake. Good historical data available. Phytoplankton biomass provides similar information to chl a concentration so choose one to avoid redundancy. |
Freshwater food web indicator evaluation
Three indicators of Freshwater Food Web community composition were identified (Table 13), but unfortunately none were evaluated for this version of the PSSU. No indicators of Freshwater Food Web energy and material flows appear on the list of candidates suggested by O’Neill et al. (2008). Indicators of this Focal Component clearly deserve greater attention in future evaluation processes.
Table 13. Summary of Freshwater Food Webs – Community Composition indicator evaluations.
Guild |
Indicator |
Primary Considerations (5) |
Data Considerations (8) |
Other Considerations (5) |
Summary Comments |
Key Fish |
Freshwater fish biomass/stream length |
Not yet evaluated |
|||
Shellfish & other inverts |
Macro invert assemblages - observed/expected |
Not yet evaluated |
|||
Key Species |
Freshwater biodiversity index |
Not yet evaluated |
Terrestrial food web indicator evaluation
O’Neill et al. (2008) identified one indicator of Terrestrial Food Web community composition (Table 14), the terrestrial biodiversity index. Unfortunately, because it is still in development, this indicator did not meet many of the evaluation criteria under the Primary, Data, and Other Considerations. No indicators of Terrestrial Food Web energy and material flows were proposed by O’Neill et al. (2008) and none were evaluated. As with Freshwater Food Webs, indicators of Terrestrial Food Webs clearly deserve greater attention in future evaluation processes.
Table 14. Summary of Terrestrial Food Webs – Community Composition indicator evaluations. The numerical value that appears under each of the considerations represents the number of evaluation criteria supported by peer-reviewed literature. For example, Terrestrial biodiversity index has peer-reviewed literature supporting 2 out of 5 Primary Considerations criteria.
Guild |
Indicator |
Primary Considerations (5) |
Data Considerations (8) |
Other Considerations (5) |
Summary Comments |
Key Species |
Terrestrial biodiversity index |
2 |
1 |
1 |
Fairly recent indicator developed by the WA Biodiversity Indicators Project. May be a good indicator for management, but more vetting required before fully usable for biodiversity assessment. |
Interface food web indicator evaluation
Two related indicators of Interface Food Web community composition were identified by O’Neill et al. (2008) (Table 15): forage fish and herring status and trends. Both indicators performed well against the Primary Considerations, though many of the Data and Other Considerations were not met. No indicators of Interface Food Web energy and material flows were proposed by O’Neill et al. (2008) and none were evaluated. In general, new, additional indicators of this Focal Component should be evaluated in the future.
Table 15. Summary of Interface Food Webs – Community Composition indicator evaluations. The numerical value that appears under each of the considerations represents the number of evaluation criteria supported by peer-reviewed literature. For example, Forage fish status & trends has peer-reviewed literature supporting 4 out of 5 Primary Considerations criteria. Details can be found in the accompanying spreadsheets.
Guild |
Indicator |
Primary Considerations (5) |
Data Considerations (8) |
Other Considerations (5) |
Summary Comments |
Key Fish |
Forage fish status & trends |
4 |
1 |
0 |
Theoretically sound and relevant, but difficult to determine whether forage fish populations are responding to management actions or pressures or environmental conditions. Highly sensitive to uncontrollable environmental conditions. |
|
Pacific herring status & trends |
4 |
1 |
0 |
Theoretically sound and relevant, but difficult to determine whether forage fish populations are responding to management actions or pressures or environmental conditions. Highly sensitive to uncontrollable environmental conditions. Good data for many Puget Sound stocks. |
About the Science Review
Puget Sound Science Review
- Ecosystem-Based Management: Understanding Future and Desired System States
- Section 1. Introduction
- Section 2. The Future of Puget Sound: Where are We Going?
- Section 3. An Approach to Selecting Ecosystem Indicators for Puget Sound
- Section 4. Evaluation of Potential Indicators for Puget Sound
- Section 5. Results of the Indicator Evaluations
- Section 6. Defining Ecosystem Reference Levels: A Case in Puget Sound
- Section 7. Glossary
- Ecosystem-Based Management: Incorporating Human Well-being
- Ecosystem-Based Management: Ecosystem Protection and Restoration Strategies
- The Biophysical Condition of Puget Sound: Biology
- The Biophysical Condition of Puget Sound: Chemistry
- The Biophysical Condition of Puget Sound: Physical Environment
- Threats: Impacts of Natural Events and Human Activities on the Ecosystem