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The provisioning of sustaining goods and services that we obtain from natural ecosystems is a strong economic justification for the conservation of biological diversity. Understanding the relationship between these goods and services and changes in the size, arrangement, and quality of natural habitats is a fundamental challenge of natural resource management. In this paper, we describe a new approach to assessing the implications of habitat loss for loss of ecosystem services by examining how the provision of different ecosystem services is dominated by species from different trophic levels. We then develop a mathematical model that illustrates how declines in habitat quality and quantity lead to sequential losses of trophic diversity. The model suggests that declines in the provisioning of services will initially be slow but will then accelerate as species from higher trophic levels are lost at faster rates. Comparison of these patterns with empirical examples of ecosystem collapse (and assembly) suggest similar patterns occur in natural systems impacted by anthropogenic change. In general, ecosystem goods and services provided by species in the upper trophic levels will be lost before those provided by species lower in the food chain. The decrease in terrestrial food chain length predicted by the model parallels that observed in the oceans following overexploitation. The large area requirements of higher trophic levels make them as susceptible to extinction as they are in marine systems where they are systematically exploited. Whereas the traditional species-area curve suggests that 50% of species are driven extinct by an order-of-magnitude decline in habitat abundance, this magnitude of loss may represent the loss of an entire trophic level and all the ecosystem services performed by the species on this trophic level.

Bhutan has measured citizens' well-being using gross national happiness since 2008 (left); GDP has been in use since the 1944 Bretton Woods meeting (right).

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Various population parameters and physiological, behavioral, morphometric, meristic, calcareous, biochemical, and cytogenetic characters have been used to identify fish stocks. We define a stock as an intraspecific group of randomly mating individuals with temporal or spatial integrity. Each character set and the associated methodology relates to specific aspects of the stock definition. Population parameters are useful primarily for the recognition of putative stocks at the practical fisheries management level. Physiological and, to some degree, behavioral characters are used primarily to study differences in the adaptation of stocks to different environments. Behavioral characters are also important for the recognition of stocks and the study of their spatial and temporal discreteness. Morphological characters, including morphometric measurements, meristic counts, and the shape, size, and type of zonation in calcareous structures provide data that are useful for the precise description of and differentiation among stocks. Although the genetic control of this type of variation is poorly understood, multivariate methods coupled with shape analyses provide techniques that describe intraspecific subdivisions that have been found to correspond to genetic stock structure as determined by other methods. Intraspecific chromosomal variation has, on occasion, been employed for stock identification. This variation has had only limited application to the study of stocks because of complications arising from intraindividual variation and artifactual variation introduced by the methodology. Electrophoresis provides an important method for measuring the genetic discreteness of stocks and for the study of genetic relationships among stocks. Electrophoretic data have recently attained a primary position among the methods used for stock identification.Key words: stock identification, genotype, phenotype, population parameters, marking, physiological, behavioral, morphometric, meristic, calcareous, cytogenetic, and biochemical characters