Modeling quantitative value of habitats for marine and estuarine populations

Coastal habitats (e.g., seagrass beds, shallow mud, and sand flats) strongly influence survival, growth, and reproduction of marine fish and invertebrate species. Many of these species have declined over the past decades, coincident with widespread degradation of coastal habitats, such that an urgent need exists to model the quantitative value of coastal habitats to their population dynamics. For exploited species, demand for habitat considerations will increase as fisheries management contends with habitat issues in stock assessments and management in general moves toward a more ecosystem-based approach. The modeling of habitat function has, to date, been done on a case-by-case basis involving diverse approaches and types of population models, which has made it difficult to generalize about methods for incorporating habitat into population models. In this review, we offer guiding concepts for how habitat effects can be incorporated in population models commonly used to simulate the population dynamics of fish and invertebrate species. Many marine species share a similar life-history strategy as long-lived adults with indeterminate growth, high fecundity, a planktonic larval form, and benthic juveniles and adults using coastal habitats. This suite of life-history traits unites the marine species across the case studies, such that the population models can be adapted for other marine species. We categorize population models based on whether they are static or dynamic representations of population status, and for dynamic, further into unstructured, age/size class structured, and individual-based. We then use examples, with an emphasis on exploited species, to illustrate how habitat has been incorporated, implicitly (correlative) and explicitly (mechanistically), into each of these categories. We describe the methods used and provide details on their implementation and utility to facilitate adaptation of the approaches for other species and systems. We anticipate that our review can serve as a stimulus for more widespread use of population models to quantify the value of coastal habitats, so that their importance can be accurately realized and to facilitate cross-species and cross-system comparisons. Quantitative evaluation of habitat effects in population dynamics will increasingly be needed for traditional stock assessments, ecosystem-based management, conservation of at-risk habitats, and recovery of overexploited stocks that rely on critical coastal habitats during their life cycle