Six
Biodiversity Research Program

Conceptual Framework

In order to describe, understand, and predict changes in marine biodiversity, well-defined, integrated research questions must be addressed. These questions must be formulated in light of the critical environmental issues prevalent in nearly all marine habitats. Furthermore, the questions must be asked across the large spatial and temporal scales defined by the open nature of most marine systems (see Box 4). The appropriate geographic and temporal scales must be explicitly defined for each regional system under consideration. Scales are in part defined by the strength of the linkages between localities, and these linkages can be determined using newly developed genetic and survey techniques (see Boxes 11 and 12) as well as through studies of the coupling of biodiversity patterns over large spatial scales (such as geographic variation in the intensity of larval settlement [Caffey, 1985; Ebert and Russell, 1988]).

Such scales tend to transcend limits normally set in biological research efforts. But traditional limits must be surmounted if the effects of anthropogenic activities on biodiversity are to be meaningfully understood in marine systems. Research tools to address biodiversity issues over the appropriate marine scales are now becoming available because of significantly improved understanding of many processes that control marine biodiversity at local scales and because of the development of new techniques for species identifications, genetic description, habitat sampling, and conducting experiments in the oceans (see Boxes 11 and 12).

For any experimental system, the questions must focus on a hierarchy of different levels: (1) patterns of biodiversity; (2) anthropogenic and natural pro-



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--> Six Biodiversity Research Program Conceptual Framework In order to describe, understand, and predict changes in marine biodiversity, well-defined, integrated research questions must be addressed. These questions must be formulated in light of the critical environmental issues prevalent in nearly all marine habitats. Furthermore, the questions must be asked across the large spatial and temporal scales defined by the open nature of most marine systems (see Box 4). The appropriate geographic and temporal scales must be explicitly defined for each regional system under consideration. Scales are in part defined by the strength of the linkages between localities, and these linkages can be determined using newly developed genetic and survey techniques (see Boxes 11 and 12) as well as through studies of the coupling of biodiversity patterns over large spatial scales (such as geographic variation in the intensity of larval settlement [Caffey, 1985; Ebert and Russell, 1988]). Such scales tend to transcend limits normally set in biological research efforts. But traditional limits must be surmounted if the effects of anthropogenic activities on biodiversity are to be meaningfully understood in marine systems. Research tools to address biodiversity issues over the appropriate marine scales are now becoming available because of significantly improved understanding of many processes that control marine biodiversity at local scales and because of the development of new techniques for species identifications, genetic description, habitat sampling, and conducting experiments in the oceans (see Boxes 11 and 12). For any experimental system, the questions must focus on a hierarchy of different levels: (1) patterns of biodiversity; (2) anthropogenic and natural pro-

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--> Box 9: Differences in environmental heterogeneity and biodiversity within and between regions provide a foundation for formulating biodiversity research questions on the patterns of diversity and why diversity matters. POLAR, PELAGIC, AND TEMPERATE-SHELF ECOSYSTEMS: CONTRASTS IN DIVERSITY AND ENVIRONMENTAL HETEROGENEITY Three regional systems—representing polar, pelagic, and temperate-shelf environments—offer tantalizing contrasts in their relative diversity and ecosystem structure. They serve as examples of the types of systems within and between which compelling research questions on the patterns of and the processes controlling diversity could be formulated: One of the most interesting contrasts in polar marine ecology is between the Antarctic and Arctic, with the former having a much higher species richness (for example, four times the number of mollusks) than the latter. Yet the Antarctic lacks the ecological (habitat and physical) diversity of the Arctic. Historical processes have led to some of the observed differences—the Arctic is heavily disturbed and has a younger fauna—but explanations for many differences in diversity and ecosystem function remain illusory. The Baltic Sea has far lower diversity than the adjacent North Sea, due primarily to differences in salinity. Despite this contrast, there appear to be few if any striking differences in energy production and flow in the water column and benthos. Both seas have similar major functional types of organisms (such as benthic macrophytes, phytoplankton, and suspension-feeding clams and mussels). What role does this level of similarity play in apparently reducing the role of diversity in energy dynamics? In the central Pacific Ocean pelagic ecosystem, the western and eastern portions appear to be comparable in species composition and structure. However, there is a significant increase in nutrient input in the eastern portion. This difference in frequency of nutrient injection may provide an excellent opportunity to determine how one scale of environmental heterogeneity does—or does not—influence biodiversity dynamics.     Key References: Elmgren (1984, 1989); McGowan and Walker (1993); Dayton et al., (1994). cesses that generate or alter these patterns, and natural processes that historically generated a given pattern; and (3) consequences to ecosystem function of biodiversity change (e.g., Box 9). Patterns Adequate knowledge of patterns of biodiversity is basic to understanding and predicting the processes responsible for these patterns. Patterns include

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--> changes in biodiversity over time and space. They include not only the species found in an area but also their relative abundances, genetic diversity, and apportionment in higher taxa. Pattern data are critical in understanding the processes that affect biodiversity and in detecting biodiversity changes. Processes Processes that determine or alter biodiversity have been studied extensively in some habitats in the sea, but generally over scales too small to allow understanding of entire marine ecosystems. Furthermore, the blending together of human impacts and natural ecological processes in pattern formation has been slow to be accepted as a theme in traditional marine ecology. Yet, successful, intensive, local studies of marine systems (e.g., the rocky intertidal; Paine, 1980; Paine and S.A. Levin, 1981) serve as critical signposts to guide regional studies of biodiversity. Moreover, retrospective studies have the potential to identify natural processes that historically generated a given pattern. These signposts point towards questions centered on critical environmental issues. When do different marine species play interchangeable ecological roles? To what extent do human activities make species or communities more susceptible to natural pressures like disease, physiological stresses, or habitat destruction? When do increasing human impacts push ecosystems across ecological thresholds? Are precipitous declines of marine species a result of these thresholds? Consequences Why does biodiversity matter? In addition to a moral and aesthetic imperative, decreasing or otherwise changing biodiversity in marine systems may have important economic effects (e.g., the collapsing fisheries around the world [Norse, 1993]). What are the most important practical reasons for maintaining natural levels of biodiversity in marine systems? This kind of question can only be answered by understanding the functional significance of biodiversity in marine ecosystems—in terms, for example, of how species diversity influences production, of how genetic diversity influences population growth or epidemics, or of how natural diversity levels confer resistance or susceptibility to invasions or to the ability of a system to recover from natural and human impacts (e.g., Tilman and Downing, 1994). These relationships are basic to understanding diversity in all environments, not just marine. In some cases, marine ecosystems provide the best experimental platforms for testing the ecological correlates of diversity in open systems. Thus, the scientific questions to be addressed would have important implications for the study and understanding of biodiversity throughout the world, providing a synergism between the unique aspects of marine and terrestrial initiatives.

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--> Basic Research Questions Listed below are some important overall questions about marine biodiversity that would constitute a major focus of this initiative. For any given marine environment, it is impossible to answer every question. However, various marine systems represent opportunities to address these questions in different combinations and ways. Fundamental advances in understanding the biodiversity of marine systems will come from coordinated studies of individual ecosystems at all three levels outlined above: pattern, process, and consequence. To apply broadly to an open system, these studies must be conducted over the scale appropriate for that system, and the results must be integrated into a dynamic view of biodiversity patterns over space and time. Because, in this initiative, the biodiversity questions would be addressed within the context of anthropogenic effects, the end result of these studies would provide critical information about practical problems in marine ecosystems, how these problems act at the ecosystem and local levels, and how connectivity between different localities may increase or buffer biodiversity change. Research conducted within this initiative is thus designed to be of a breadth, depth, and scale such that a more complete environmental picture can emerge, facilitating many potential management and conservation applications. Natural Variation in Biodiversity Pattern and Why Biodiversity Matters: How do genetic, species, and ecosystem diversity vary in space and time at different regional-scales, and within habitats within these regions? Examples of specific research questions are: To what extent does the maintenance of local biodiversity (genetic or species) depend on linkages between distant populations, the dispersal between them, and the availability of suitable habitat? How does genetic diversity within a species influence reproduction and population growth or susceptibility to epidemic disease? To what extent do changes in biodiversity at one site within a region—or between regions—affect the biodiversity at another site or in another region? What specific characteristics of a habitat directly or indirectly influence genetic and species diversity? For example, are there parallels in the origin and maintenance of coral reef and deep-sea biodiversity? What is the functional significance of biodiversity at the genetic, species, and ecosystem levels? Are species within a functional group interchangeable? What might be learned from comparing and contrasting systems in terms of the functional significance of biodiversity? (For example, are there parallels between the ecological significance of microbial diversity as coral reef symbionts [zooxanthellae] and as open-ocean primary producers [picoplankton]?)

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--> To what extent does the diversity of a community determine (a) "stability," (b) productivity, (c) resistance to invasion or disease, and (d) ability to recover from natural and human impacts? Equally important, how do these factors interact? Do high diversity systems have higher or lower production than systems whose diversity has been impaired? What is the role of biological invasions in altering system production or energy flow? How good are the estimates of genetic, species, and ecosystem biodiversity, and how do the limitations (i.e., understanding of the scale of error) influence an understanding of biodiversity patterns and of ecosystem structure and function? Human Impact on Processes Responsible for Biodiversity Change: What are the direct impacts on biodiversity of human-altered systems? That is, what is the variation in biodiversity over spatial and temporal scales relevant to the critical environmental issues? Examples of specific research questions are: How do human influences on biodiversity differ from those caused by natural processes? To what extent do human effects alter the probability of ecosystem collapse in different systems? To what extent are particular changes in biodiversity due to human activities reversible? Given the often direct impacts on certain target species within a region, are species within functional groups interchangeable within a system? How does the addition or loss of species due to human activities affect community structure and resilience? What are the indirect impacts on biodiversity of human-altered systems? Examples of specific research questions are: What characteristics of species enhance susceptibility or provide immunity to precipitous declines? In what types of habitats are alternative ecological communities stable? Are threshold processes involved in precipitous declines (and the persistence of these declines) in biodiversity, and, ultimately, in the risk of extinction of individual species? Does genetic or species diversity provide a buffer against irreversible or massive perturbations? What are the long-term effects of species replacements (e.g., exotic species) on ecosystem function?

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--> Contingent and Reasonable Prediction In this proposed research program, where a major goal is prediction of the effects of human activities on marine biodiversity as input to conservation and management decisions, it is important to recognize and characterize the uncertainty associated with such predictions. Prediction stems from a basic understanding of system processes, and this initiative is designed toward this end. Moreover, this initiative emphasizes the need to expand the predictive capability of the marine sciences within a framework contingent on appropriate spatial and temporal limitations, as well as on limitations in other aspects of the science. Such limitations should be expressly considered in developing predictions. Mechanistic models are valuable tools for predicting and extrapolating beyond the specific parameter range studied, and sensitivity analyses for terms in the models can provide at least qualitative estimates of the uncertainty associated with such predictions and extrapolations. In the ocean, no diversity inventory can ever be complete, and due to the vastness of even regional sites, large patterns must be discerned with proportionately few samples. Therefore, inference must be an important tool in the study of marine biodiversity. Thus, in order to assure the efficiency of research and the rigor of predictions, it will be necessary to understand fully, to develop, and to advance strong inferential methods (e.g., Platt, 1964). In addition, the consequences of taxonomic error are not typically addressed in ecological studies, but appreciation of such consequences is critical to evaluating the reliability of the biodiversity patterns that would form the foundation of the research conducted in this program. Thus, the taxonomic component of this program includes formal evaluation of the consequences of taxonomic error and changing taxonomic resolution on the design, analysis, and inferences drawn from the biodiversity studies (Carney, 1993; Carney, in press). Approaches Theory and Modeling The central biodiversity research questions require the means to describe and characterize patterns of biodiversity, an understanding of biodiversity for ecosystem processes, and an understanding of the mechanisms that maintain biodiversity. Each of these challenges involves relating patterns and processes across multiple scales of space, time, and organizational complexity and mandates the use of modern approaches to modeling and computation. Which details at fine scales are important to understanding patterns on broader scales, and which are irrelevant? How can the dynamics of aggregations—populations, communities, or other groupings—be understood in terms of the collective motions of the individuals? How much functional redundancy exists in ecosystems, and how much functional detail is needed to account for shifts in ecosystem response?

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--> Each of these questions involves a recognition that systems are structured into components that have characteristic spatial and temporal scales, and that only macroscopic descriptions of the dynamics within components may be relevant to the dynamics of the collections of components. But to determine the identity of those macroscopic descriptors requires a melding of bottom-up, individual-based approaches and top-down phenomenological approaches. Only in this way can the mechanistic basis of the patterns observed be explored, for example, in remote sensing and other broad-scale approaches to description. This also enables the broad-scale predictions of climate models to be interfaced with the much finer-scale understanding that exists at the level of the individual. The primary frameworks for describing the dynamic movements of populations and communities are those associated with the concepts of Lagrangian and Eulerian motion. In the former, individuals (or packets of fluid) are traced as they make their way across a landscape or seascape. In the latter, particular regions of space provide the starting point, and numbers of individuals (or the densities of materials) in the region are tracked, describing changes in terms of fluxes; individual identities are suppressed. The two approaches provide different perspectives on the dynamics of population and communities. The study of biodiversity must utilize both modeling approaches. Individual responses to changing environmental cues, and to each other, embody knowledge of organismal-level behavior and physiology and allow the representation of taxonomically based functional biodiversity. But observations are most easily expressed in Eulerian terms, reflecting patterns, for example, of the type detected by remote sensing. Hence, exciting and powerful new advances in modeling will involve the development of individual-based models that allow prediction at the ensemble (aggregate) level, or in an Eulerian frame of reference, and techniques for relating Lagrangian and Eulerian models (e.g., Grunbaum, 1992; Grunbaum and Okubo, 1994). Great advances have been made in numerical modeling of the oceans in the past few decades. Models can be used to predict the variability in physical properties of the seawater, the general circulation of the oceans, and small-scale changes in flow patterns that might result from anthropogenic causes (such as the building of groins, etc.). Of particular concern within the context of this initiative is the incorporation of biology into physical models. Until recently, organisms have been treated as passive (i.e., nonswimming, noninteractive) tracers in the models. Ideally, dispersal and recruitment can be predicted, given a larval source. In addition to all the usual uncertainties associated with numerical modeling of geophysical fluid dynamics, some of the physical limitations of these models that are particularly relevant to this program include the uncertainty in the diffusivity coefficients and the poor resolution of vertical motions, especially on short time scales. Over the last decade or so, much progress has taken place in the development of coupled models of biological, chemical, and physical systems. This has been

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--> and is being done both on a global scale using "gross" processes (Fasham et al., 1993) and on smaller-scales with more detailed interactions (Hofmann, 1988; Hofmann and Ambler, 1988; Werner et al., 1993; Tremblay et al., 1994). In the coming years, the incorporation of spatial dependence into individual-based modeling (DeAngelis and Gross, 1992) is sure to advance this field significantly. The utility of numerical models for understanding biodiversity phenomena within an oceanographic setting would be improved considerably by the incorporation of effects such as aggregation or behavioral mechanisms (including swimming and vertical migration) that would tend to clump or separate organisms (e.g., Rothlisberg et al., 1983; Franks, 1992; Eckman et al., 1994). These types of models would be particularly useful in addressing connectivity between sites within a region via larval dispersal and settlement (in the case of planktonic propagules of benthic organisms). Ecological modeling will also be valuable for designing experiments, interpreting results, and making predictions regarding, for example, the role of life-history characteristics in determining the susceptibility of species to global crashes (leading toward extinction), and the probability of subsequent recovery. In such metapopulation models (Nee and May, 1992), the following four crucial factors are involved in predicting, for a given species, the threshold amount of habitat that can be lost and the distribution of remaining habitat for sustaining the species: (1) the total numbers of semi-isolated populations (patches) of competitively superior and weedy species, (2) the health of those populations (e.g., are they increasing, decreasing, or extinct), (3) the disturbance rate that would obliterate local populations/patches (e.g., epidemics, hurricanes), and (4) the colonization abilities of different species that allow them to locate new patches, which are a function of the species' life-history characteristics and the physical connectivity between different patches. Such models should be an integral part of this initiative because they explicitly relate the local demographic and life-history characteristics of a species and its most important neighbors, and the physics of their "regional-scale" habitat (see also Mangel, 1993). These models also indicate the clear need for both smaller-scale, local studies of the population biology and ecology of species, and of a larger-scale perspective concerning the physical connectivity between suitable habitat, and thus populations. These models dictate the most critical biological and physical parameters that should be measured or manipulated in the field. Moreover, they can be used to develop management criteria for protected areas, particularly because the model output includes predictions of the minimum number of patches required to sustain and "protect" the species. Retrospective Analysis: Importance of an Historical Perspective Recent history provides a valuable yet underutilized guide to the factors controlling the number of species occurring at each of several spatial scales. It

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--> also enables estimates of rates of change in species numbers and thus provides an important scale against which to measure future changes (Elmgren, 1984, 1989; Webb and Bartlein, 1992; Ricklefs and Schluter, 1993). Differences in species richness among geographical regions of the modern ocean are, to a large extent, attributable to differences in the magnitudes of extinction, diversification, and immigration of species during the last 20 million years of Earth history. Studies of geographical range contractions reveal not only which factors might be responsible for local and global extinctions, but more importantly, which characteristics of habitats and regions enable species to persist during crises. Similarly, comparisons of fossil biotas with living ones from the same habitats and regions provide clues about conditions that are favorable to the formation and proliferation of species. Such comparisons are especially meaningful and powerful when they are done in conjunction with phylogenetic analyses, because the latter make it possible to trace ancestor-descendant relationships and provide one means by which to infer when, where, and how frequently new species arise. Finally, the fossil record offers both a uniquely long-term perspective on biotic interchange—the movement of species across geographic barriers that have partially or completely broken down (Vermeij, 1987, 1991a, 1991b)—and on the temporal pattern of diversity and rates of origin and extinction of marine species (Jackson et al., 1993; Jackson, 1994). Given that such interchange potentially affects diversity and that it is taking place through human agency at unprecedented rates, it is important to investigate the extent and consequences of historical cases of biotic interchanges at scales ranging from particular habitats to biogeographic provinces. These studies are most profitably done with taxonomically well-characterized groups having a good fossil record. Foraminiferans, mollusks, corals, sea urchins, barnacles, bryozoans, marine birds and mammals, diatoms, and calcareous algae would thus be particularly promising groups for study. Another promising historical approach that has been almost entirely ignored is the systematic study and evaluation of museum or archived collections (Box 10) and of monographs from the eighteenth through the early twentieth centuries. Buried in these early collections and papers are important records of species whose ranges have contracted or expanded since then, or which have been wholly exterminated. An example is the logbooks of whalers in the nineteenth century, whose observations of whale populations form the basis of estimates of preexploitation population sizes (Evans, 1987; Baker and Palumbi, 1994) and whose pages may literally provide estimates of genetic diversity in previous centuries. From such material it may be possible to estimate rates and times to extinction, rates and times of species introductions and invasions, and other changes in geographic distributions of common species during the last two centuries. This historical research would profitably include in-depth examination of documents related to shipping and fisheries, as well as the proceedings of local scientific societies and of groups devoted to natural history.

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--> Box 10: Museum collections of biological specimens are an invaluable and largely underutilized resource for biodiversity studies. THE UTILITY OF MUSEUM COLLECTIONS Some of the first attempts to describe the diversity of species at the base of the food web were made by fisheries biologists in the 1870s and 1880s. These collections, made by biologists aboard the U.S. Fisheries Steamers Fish Hawk, Albatross, and Blake, can be found today on shelves deep in the recesses of the Smithsonian Institution's National Museum of Natural History. Many tens of millions of other specimens from the waters of the United States and around the world are preserved in that and other museums in this country. These specimens represent a partial catalogue of the Earth's natural history, and, for those species that are described, provide the standards for names of the world's species. But these specimens can also be used to document biotic changes in areas of habitat alteration, to analyze long-term changes in species distributions, and to determine both the historical and modern importance of the incorporation of human pollutants into marine food webs. Thus, museum specimens of tuna (Thunnus spp.) were analyzed to determine the historical trends of mercury contamination. In addition, museum collections provide a major potential resource for determining species extinctions of both animals and plants in the ocean. This approach could include: (1) locating specimens of species (especially from well-collected coastal habitats) that have not been found again in the twentieth century, and (2) re-examining, through molecular genetic analysis, preserved specimens of extinct populations of species that have been thought to be conspecific with extant populations.     Key References: Lee et al., (1978); Carlton (1993); SA2000 (1994). Archived, long time-series data are also invaluable for retrospective studies. Long-term sets of physical, chemical, and biological data exist that can provide historical records up to about one century old. The more than five decades of data from the California Cooperative Ocean Fisheries Investigations (CalCOFI) program is one example (Chelton, 1983). Historical accounts and museum collections (Box 10; Jackson, 1994) provide resolution extending to several centuries. Archived cores from individual corals, analogous to tree ring cores, can yield valuable historical information on reef history, El Niño-Southern Oscillation events, and other climatic episodes, for up to perhaps five centuries (Glynn and Colgan, 1992; Dunbar and Cole, 1993). Finally, archived sediment cores and cores of entire coral reefs provide historical environmental, biological, and geological data ranging from thousands of years (with resolution at the level of decades) to hundreds of thousands or several million years (with resolution ranging from thousands to tens of thousands of years) (Jackson, 1992; Webb and

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--> Bartlein, 1992; Valentine and Jablonski, 1993). The use of such existing samples or data sets will allow examination of temporal and spatial scales precluded by short-lived grants, and may also prove to be less expensive than new field studies. In summary, the historical approaches outlined here offer a perspective on marine biological diversity at longer time scales. They place taxonomic, ecological, and biogeographical studies in a larger framework in which the appearance and disappearance of species can be linked to known mechanisms and events of environmental change. Linking Biotic Surveys with Ecological and Oceanographic Experiments Within a regional system, taxonomic surveys of those taxa fundamental to pattern-level and process-level questions should be done as an integral part of the research program. In this sense, biotic surveys are vitally linked to the ecological and oceanographic regional-scale approach identified here. Many biotic surveys currently exist through fisheries or agency-based activities. Fishery research vessel cruises, for example, sample the ocean widely but typically target a relatively narrow range of species; with additional effort these surveys could be expanded to include more comprehensive biological sampling. Similarly, sampling from fishing vessels, many of which have scientific observers on board, would be an additional source of information. Marine fisheries also represent one of the greatest manipulations of marine ecosystems by the human race. Management approaches take into account not only biological, but also social, economic, and international considerations. Although not widely applied, the approach of adaptive fisheries management, where fish populations are manipulated to learn about the processes regulating their population sizes (Walters and Hilborn, 1978; Collie, 1991), would provide excellent opportunities for studying attendant biodiversity-related issues within the context of this initiative. In a similar manner, some research under this initiative should take advantage of fishery management regimes to examine ecosystem response carefully and to monitor, and ultimately predict, the consequences in terms of biodiversity. Methods Studying the effects of environmental change and anthropogenic activities on regional-scale marine biodiversity and the consequences to ecosystem function presents unique conceptual and methodological challenges. Much previous ecological work has focused on relatively circumscribed spatial/temporal scales that do not adequately account for the vast geographic ranges of many species. Conversely, relevant spatial scales for smaller organisms and microorganisms have often been inadequately described and undersampled. In addition, although

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--> applied within and across diverse sets of taxa. Molecular methods are now an important component in many biodiversity research programs. Integration and confirmation of such techniques with traditional taxonomic criteria is a necessary and important component of emerging biodiversity research programs worldwide (NERC, 1992). Instrumentation In situ field approaches and innovative instrumentation are having large impacts on marine biodiversity research. A good example is the application of flow cytometric techniques in the recent discoveries of phytoplankton diversity (R.J. Olson et al., 1991). Newly recognized prochlorophytes in the photosynthetic picoplankton were discovered in oligotrophic ocean gyres (see Figure 6) by recognition of their unique pigment signatures using shipboard flow cytometry (Chisholm et al., 1988). These abundant planktonic organisms were largely unnoticed until application of this new technology. This example illustrates the profit of welding different approaches and technologies. In subsequent molecular studies (Giovanonni et al., 1990; Schmidt et al., 1991), an abundant ribosomal RNA sequence, peripherally related to marine Synechococcus sp., was identified in bacterioplankton assemblages from oligotrophic ocean gyres. Subsequent genetic analysis of cultured marine prochlorophytes showed that this sequence was derived from the very same prochlorophytes previously identified by flow cytometry. Use of tagged molecular markers such as fluor-labeled oligonucleotides or antibodies, in combination with sensitive instruments such as flow cytometers, have great potential for assays of biodiversity in natural and anthropogenically altered systems (Ward and Carlucci, 1985; DeLong et al., 1989; Amman et al., 1990). Sampling Improvements in sampling methods and in situ approaches will be useful for regional-scale studies of biodiversity (Box 12). For pelagic studies, in situ plankton samplers and fluorimeter devices are being developed for use in tandem with sensitive molecular identification methods for detecting and quantifying the presence and variability of specific larval and microbial taxa. Real-time surveys of variability and abundance of microbial and metazoan organisms are now entering the realm of possibility by coupling sophisticated electronics with molecular identification techniques, but more development will be needed for widescale application of such technology. Ribosomal RNA probes already have been used in conjunction with flow cytometry in analyzing mixed microbial populations (Amman et al., 1990). In situ field sampling and detection represents an important, achievable advance in broad-scale marine biodiversity studies. Remote

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--> Box 12: Conducting regional-scale research will require the application of many new sampling and observing technologies. THE ROLE OF NEW SAMPLING AND OBSERVING TECHNOLOGIES Techniques long used in terrestrial studies of biodiversity are now being extended to marine systems. Remote-sensing technology provides a critical tool to evaluate linkages between ecological and oceanographic processes in regional-scale studies. Satellite information can now be used to determine primary production and physical transport processes that influence biodiversity. Navigational capabilities are now accurate to one centimeter, and geographical information systems (GIS) allow storage of images and data at a variety of spatial scales. These advances revolutionize the ability to analyze long-term biological and physical-oceanographic data on many temporal and spatial scales in conjunction, for example, with data on climate changes. In turn, marine scientists must be closely involved in the design and application of satellite sensors. In sublittoral habitats, obtaining information for the study of biodiversity has been more difficult. Now, however, 10-meter contour topographic maps are available for some sites and centimeter-scale maps are possible. Bottom images from high-resolution side-scan sonar can provide an intermediate-scale reference so that photographic images can be overlaid precisely on multibeam sonar topographic maps. These recently developed techniques can begin to provide a spatial information reference taken for granted in terrestrial studies. Low-cost remotely operated or autonomous vehicles can be launched from shore stations to study the horizontal extent of events observed either from satellite imagery or in situ observing systems. In order to study adequately processes controlling biodiversity, continuous data in both space and time are needed to record low-frequency events. The resultant maps and time-series are needed to guide sampling and to design experiments. Developments in the assimilation of data using nested series of models will enable analysis of the vast amounts of spatially continuous, real-time data generated by ocean-observing systems (in moorings, bottom landers, and via electro-optical cable connections to land). Awareness of the ways in which different organisms interact with oceanic processes is only just beginning. There are new optical sensors for every scale of resolution from satellite-captured views from space to in situ microscopes; acoustic sensors developed for detecting submarines can be used to study whales, and other acoustic instruments are being used to study fish and their microscopic food. Moreover, it is now possible to visualize the physical, chemical, and biological surroundings of individual planktonic larvae or zooplankton. These first-hand views of biological as well as physical patterns and structure in the ocean will lead to better scientific questions, better experiments, and better management of marine systems.     Key References: Greene et al., (1988); Currin et al., (1990); Greene and Wiebe (1990); Abbott and Chelton (1991); GLOBEC (1991, 1992); Von Alt and J. F. Grassle (1992); USGS (1994).

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--> sensing and acoustic techniques are also important methods for providing valuable information on regional-scale phenomena (Pieper and Van Holliday, 1984; Greene et al., 1989; P.E. Smith et al., 1989; Fua et al., 1990; Abbott and Chelton, 1991). Although these techniques may not have sufficient resolution to provide specific information on species or intraspecific biodiversity, they can provide broad-scale perspective on bulk environmental properties or processes that may influence, or be affected by, changing patterns in biodiversity. Moreover, high spectral resolution imagery, for example, may be able to distinguish different phytoplankton pigment groups using satellite-based sensors. Carder et al., (1993) used an aircraft-based sensor to study the distribution of substances other than chlorophyll in the coastal ocean. Broad-scale changes in pigment composition may be indicative of changes in the physical environment (Letelier et al., 1993). New, automated, direct-sampling technology, such as the Moored, Automated, Serial Zooplankton Pump (MASZP) (Doherty and Butman, 1990; Butman, 1994) and new optical imaging technologies such as the Video Plankton Recorder (VPR) for zooplankton (Davis et al., 1992), the Rapid Sampling Vertical Profiler (RSVP) for phytoplankton (Cowles and Desiderio, 1993), and the EcoScope (Kills, 1992) for studying predator-prey interactions show great promise for providing species-specific distributions of planktonic organisms over relatively large geographic regions. Some of the best opportunities for technological advances in biological oceanography involve coupling automated, direct-sampling methods with automated, sample-processing techniques. Work is in progress (C.A. Butman and E.D. Garland, pets. comm., 1994), for example, to develop species-specific, fluor-tagged immunological markers for planktonic larvae of benthic invertebrates that can be applied to samples collected by the MASZP. Then, the number of fluorescing organisms can be quantified automatically using digital, color image-analysis techniques (Bjørnsen, 1986; Amman et al., 1990; Berman, 1990; Sieracki and Viles, 1990). This is a good example of the enriching opportunities to couple existing technologies developed in diverse fields of science and engineering for new applications in biological oceanography and marine ecology. Special Opportunities Coordination with "special opportunities" in environmentally relevant marine biodiversity research should be recognized. In particular, recognition of threatened species in fisheries or inventory studies, or identification of invasive species may provide unique "samples of opportunity" in marine biodiversity research. One example is coordinated studies in collaboration with fisheries experts on recently impacted or endangered species or populations. Other coordinated studies on biodiversity could readily take advantage of long-term, process-oriented biogeochemical time-series studies already in place. Such opportunities should not substitute or supplant, but rather augment, the organism-oriented

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--> biodiversity emphasis of this initiative. Finally, the establishment of marine reserves offers one of the most striking opportunities for understanding human impacts in the sea (Box 13). It is clear that understanding of the distribution of marine organisms, and therefore their conservation and management, will require studies at unprecedented geographic scales. The marine laboratories of the world have great potential to provide the infrastructure and focus for programs in research, training, and education, and the conservation of marine biodiversity. Regional marine laboratories encompass the geographic scale of environmental and ecological gradients and bridge the disciplines of oceanography and ecology, and their region-wide data sets are fundamental to structuring comparative studies of marine biodiversity (Lasserre et al., 1994). Marine laboratories are found in virtually every coastal country, often in relatively undisturbed locations, with ready access to many representative coastal habitats and organisms. The great majority of laboratories are tied to academic institutions or museums, with long-standing traditions in the study of marine organisms, training of scientists and managers, communication and exchange with other laboratories, and environmental impact assessment. Many marine laboratories are government-supported, with strong mandates for resource management. Their continuity of research and management sets marine laboratories apart from other institutions. They either possess or have direct access to unique, long-term data sets that form a critical baseline against which human impact may be assessed and interpreted. Whereas marine laboratories are found within different countries or regions with different cultures, they have a common scientific culture and traditions which predispose them to cooperative programs and to networking. For example, the 27 laboratories of the Association of Marine Laboratories of the Caribbean (AMLC) have held annual meetings for almost 30 years. In 1990, with the support of the National Science Foundation and private foundations, over 20 Caribbean laboratories formed the CARICOMP (Caribbean Coastal Marine Productivity) network to conduct comparative, standardized observations of coastal ecosystem structure and function (Ogden, 1987). More recently, 80 European laboratories have joined together in the Marine Research Stations Network (MARS), and U.S. marine laboratories have formed the National Association of Marine Laboratories (NAML), as well as regional groups such as the 35-member Southern Association of Marine Laboratories (SAML) (Lasserre et al., 1994). Implementation In order for this national research agenda to be realized, the scientific community, federal agencies, and key related programs will need to work together in a coordinated, committed fashion, tapping into and building on the mounting enthusiasm for tackling the challenges of the critical environmental issues now facing the oceans. A consequence of this realization will be that the "payoffs"

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--> Box 13: Marine research reserves provide many special opportunities for addressing biodiversity research questions in controlled and protected settings. MARINE BIODIVERSITY RESERVES FOR RESEARCH, CONSERVATION, AND MANAGEMENT Marine reserves are essential for measuring human impacts on the diversity and stability of communities and for developing more scientifically rigorous guidelines for their protection and management. Whereas manipulative experiments to alter the abundance of selected species or to evaluate environmental changes under controlled conditions are the best way to address many important biodiversity questions, experiments have rarely been conducted at the large-scales appropriate to understanding why communities change. Marine reserves would be excellent for addressing questions such as the following: What are the physical effects of human activities on coastal biodiversity? Extraordinary effects were demonstrated, for example, when people were excluded from rocky intertidal shores of Chile, resulting in nearly total transformations of intertidal communities in only a few years. How large an area is required to protect nursery populations that are fished elsewhere, and over how large an area might such nurseries be effective? Chilean workers have demonstrated that some marine reserves do not work well in supplying adjacent regions with larvae of overharvested species. How well can degraded environments and communities be restored, and how long does it take? How effective is the re-establishment of endangered species into reserves? Sea otters, for example, have been re-established successfully in their native habitat in the Pacific Northwest. There are many excellent opportunities to conduct large-scale human exclusion experiments within existing parks, sanctuaries, military bases, and other areas where public access is already restricted. Areas closed due to collapse of fisheries are also good prospects. The number, size, and treatment of reserves need to be carefully planned in advance to take advantage of regional knowledge and for statistical analyses. This is critical for experiments investigating the potential for multiple-use reserves. Finally, human-exclusion experiments provide an opportunity for education about important marine resources to facilitate public support for hard management decisions, particularly those involving recreational activities. Support for such restrictive measures could be increased by involving local residents in experiments.     Key References: VanBlaricom and Estes (1987); Duran and Castilla (1989); D'Elia et al., (1991); Siegfried (1994).

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--> and products that can result from this first national marine biodiversity agenda will contribute significantly to the nation's emerging agenda on the conservation and wise use of the nation's biological resources. This initiative will require major new funding over a decadal time scale to achieve the objectives outlined in this report. In addition, existing programs and resources from federal, state, and private sources would augment any new funding (e.g., see Box 13). We address here the broad aspects of implementation and directions for this marine biodiversity initiative. Future study plans will develop the specific processes and mechanisms for accomplishing the objectives of this initiative. This proposed initiative would be propelled by the interactions and involvement of several agencies working under coordinated research umbrellas. Energetic and intimate working relationships with national and international biodiversity programs, marine laboratory networks, museums, and newly emerging fields (e.g., Box 14) will form critical research bridges. This initiative is envisioned in terms of a decadal time scale—a minimum time period to undertake the research efforts proposed here and to achieve a critical level of coordination. The research questions posed in this agenda are envisioned as being addressed by both small and large research groups seeking designated funds from several agencies through a peer-review process. Once launched, early steps would include regional integration of research efforts. Coordination may include specific efforts to use the same experimental techniques and sampling methods to address the same or similar questions across different systems. The selection of which systems to study in which geographic regions should ultimately be determined by the competitive proposal process. Important decision criteria for funding would include the extent to which the proposed research addressed scientifically perceived environmental threats to the identified study system and the likelihood of achieving substantial new insights that can be applied to conservation and management. The regional-model system approach has the virtue of concentrating effort, but could risk too great a focus on special cases that may be so exceptional as to limit future applications and generalizations. To minimize such biases, regional-model systems should be chosen as much for their diversity of characteristics as for their taxonomic variety and imperilment. This marine biodiversity initiative is not calling solely for entirely new data collections, experiments, and investigations. The existence of critical, and especially, long-term information or archived samples for a given site, region, taxon, and so forth, will be invaluable for interpreting results of new studies undertaken as part of this program. Such data would decidedly leverage time, effort, and resources spent on conducting new studies. In some special cases, existing data sets (for example, the CalCOFI data mentioned earlier) may be sufficient for addressing a given suite of biodiversity research questions. As noted by Angel (1992), "The information base on which to develop a predictive understanding of the interaction between diversity and ecological process can be greatly enhanced relatively inexpensively by systematically collating existing data and working up extant collections of material."

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--> Box 14: Knowledge resulting from this research program should have important economic impacts, for example in the newly emerging enterprise of marine biotechnology. MARINE BIOTECHNOLOGY AND MARINE BIODIVERSITY: ARE WE LOSING RESOURCES BEFORE THEY ARE EVEN RECOGNIZED? The newly emerging and rapidly developing field of marine biotechnology depends for its very existence on a constantly new and broadening knowledge of resource organisms for biomedical, bioengineering, and mariculture research. The potential application of information that could be derived from this initiative for the biotechnological frontier includes natural products derived from marine animals and plants in all ocean habitats, preservation of marine genomic information, development of hardy culture stocks, resistance to disease, and closed-system, computer-controlled aquaculture of marine species. These are but a few examples of the total potential waiting to be tapped. Loss of biodiversity in tropical terrestrial ecosystems, long before it is even recorded, and the concomitant loss to the biomedical and biotechnological sciences, are now well documented. Loss of biodiversity in the world's oceans (especially given the much greater higher-order diversity in the oceans, and thus the potential loss of the genome and chemical makeup of an entire class or order of organisms), and its potential value for solving pressing medical and food problems of the growing human population would be one of the great tragedies of our time.     Key References: Colwell (1983); Fautin (1988); Marine Life Resources Workshop (1989); Colwell and Hill (1992); Weber (1993). Relationship to Other Programs "An apothegm applies here: a person with one watch knows what time it is; a person with two watches is never sure. Those who espouse solely the approach based on carbon stocks and flows, or solely the approach based on food webs, can confidently contrast marine and terrestrial biotas. Those who consider both approaches have grounds for further thought". Cohen (1994, p.63) This initiative proposes research on regional-scales not previously undertaken in the marine environment. The decision to work at these larger scales suggests fundamental and potential linkages and synergisms with other programs that have similar or overlapping interests. In addition, the missions of many federal agencies increasingly call for addressing critical marine biodiversity is-

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--> sues. This initiative provides the next step to connect and coordinate research opportunities with other programs and with agency interests and efforts. In turn, oceanographic-ecological coupling within a regional-model system approach means that this initiative could provide numerous potential benefits for other biodiversity programs. Scientific Programs and Initiatives The Systematics Agenda 2000 (SA2000, 1994) is a consortium-based program that seeks to improve the discipline of systematics and taxonomy throughout the U.S. scientific structure. The importance of accurate and reliable taxonomy to studies of biodiversity cannot be overemphasized. Thus, the goals of SA2000 coincide with some of the goals of this initiative—enhancing taxonomy as a discipline and improving the scientific knowledge base of the systematics of marine organisms. This marine biodiversity initiative further recommends an enhanced interrelationship between taxonomists and ecologists. Systematic support groups, such as the Association of Systematics Collections (ASC), have formulated plans for the conservation, support, and use of the nation's museum collections. The Diversitas program, sponsored by the International Union of Biological Sciences (IUBS), the Scientific Committee on Problems of the Environment (SCOPE), the International Union of Microbiological Scientists (IUMS), and the United Nations Educational, Scientific and Cultural Organization (UNESCO) is a broadly based program in biodiversity covering all of the Earth's ecosystems (diCastri and Younès, 1990; 1994). Under the auspices of the International Council of Scientific Unions (ICSU), Diversitas has become one of the four components (along with the World Climate Research Program, the International Geosphere-Biosphere Program, and the Human Dimensions of Global Environmental Change Program) of a comprehensive program of Earth systems research (Perry, 1993). The scientific questions asked by Diversitas are closely related to those in this initiative. The marine component of Diversitas is described by J.F. Grassle et al., (1991); the European MARS Network, noted earlier, is a component of Diversitas. Diversitas has sponsored workshops on biodiversity and ecosystem function for many habitats, including coral reefs; upwelling systems; estuaries, lagoons, and nearshore coastal systems; and pelagic systems. These workshop reports and conclusions are a rich source of research questions closely related to those identified here. This marine biodiversity initiative would benefit—and would benefit from—some of the other major marine initiatives such as the Joint Global Ocean Flux Study (JGOFS) and the Global Ocean Ecosystem Dynamics (GLOBEC) program. One of the objectives of JGOFS is to ''understand the global-scale processes that control carbon, nitrogen, oxygen, phosphorus, and sulfur exchanges in the ocean over time" (NRC, 1994b). The goal of GLOBEC is "to predict the

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--> effects of changes in the global environment on the abundance, variation in abundance, and production of marine animals," with an emphasis on how changing climate alters the ocean's physical environment and how such alterations affect zooplankton and fish (NRC, 1994b). This proposed initiative has clear synergistic linkages with programs like JGOFS and GLOBEC; research questions called for here provide a framework, for example, for addressing the role of biodiversity (particularly at the microbial level) in ocean carbon flux, and thus the precise biological pathways through which carbon may pass in the ocean. Similarly, this initiative identifies global climate change as a critical environmental issue relative to potential changes to the oceans' biodiversity, and thus this initiative forms a natural linkage with other efforts that seek to understand the biological and oceanographic mechanisms that may influence the temporal and spatial variation of populations in the sea. Federal Agency Programs Numerous federal programs address a wide variety of biodiversity technologies and issues. Programs within the Departments of Commerce, Interior, Energy, and Defense, the National Institutes of Health (NIH), the National Science Foundation (NSF), and the National Science and Technology Council (NSTC), focusing on such aspects as genome mapping, new information technologies, bioremediation, and environmental quality assessment, offer important opportunities for interaction with many aspects of this proposed marine biodiversity initiative, as well as for the cooperative use of resources. Within NSF, research on marine biological diversity is funded within the Ocean Sciences Division (Directorate for Geosciences), the Office of Polar Programs, and the Biological Sciences Directorate. The first two offices currently focus primarily on basic science issues relating to: ecological, evolutionary, and historical processes responsible for maintaining or changing diversity in a system; the functional role of diversity in ecosystems and populations, and the importance of species/gene redundancy; the impact of introduced species on natural ecosystems; developing advanced procedures for resolving taxa in traditionally recalcitrant groups (e.g., microbes, algae, planktonic invertebrates); characterization of diversity in marine systems, including coastal, open-ocean, sea ice, and deep-sea systems; and biogeography related to dispersal processes. These offices are also concerned about the declining expertise in taxonomy and systematics, and the ongoing maintenance of living collections and laboratory facilities for research in marine biological diversity. NSF's Biological Sciences Directorate (primarily through the Division of Environmental Biology, but including the Divisions of Molecular and Cellular Biosciences, and of Integrative Biology and Neurosciences) currently supports some marine biodiversity research activities as part of its general funding of biodiversity, including Biotic Survey and Inventory Research, Basic Research in

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--> Conservation and Restoration Biology, Partnerships for Enhancing Expertise in Taxonomy, and the Joint International Research program administered cooperatively with the U.S. Agency for International Development (AID). The National Oceanic and Atmospheric Administration (NOAA), in the Department of Commerce, has a diverse mission that includes promoting environmental stewardship in order to conserve and manage the nation's marine and coastal resources. NOAA has a long history of work in marine biological diversity, stemming from over a century of physical and biological surveys and two decades as the federal trustee for the conservation, protection, and management of fishery resources, marine mammals, and endangered marine species. NOAA is also responsible for managing Marine Sanctuaries and Estuarine Research Reserves, and has the federal mandate to evaluate and restore marine ecosystems impacted by oil spills and other human disturbances. The 1992 NOAA Strategic Plan (NOAA, 1992) states that the inventory, assessment, and maintenance of marine and coastal biodiversity is an important goal of the element entitled Environmental Stewardship: Coastal Ecosystem Health. New opportunities for large-scale experimental approaches would complement NOAA's efforts to manage and protect fisheries and other species, as well as their habitats. NOAA's National Marine Sanctuary Program and the National Estuarine Research Reserve system have programs to inventory biodiversity and begin the monitoring and assessment of long-term trends within these protected areas. These programs provide yet another cooperative platform for research. As the primary agency concerned with marine biodiversity within the national partnership for biological survey (of which the National Biological Service [NBS], described below, is a part), NOAA's mission and programs align nicely with the research framework provided by this marine biodiversity initiative. The Office of Naval Research (ONR), within the Department of Defense, relies on continual improvements in the understanding of marine biodiversity and its impacts on natural marine communities. As a steward of the marine environment, ONR programs target research that contributes to preservation and conservation of marine resources and diversity, and to the development of technologies that eliminate or minimize potentially harmful impacts of naval operations on the marine environment. A major thrust area is the development of biotechnical capabilities that exploit the rich diversity of marine organisms to provide new materials, processes, and capabilities. ONR's programs also target understanding of the ocean biota in relation to the chemistry and physics of the sea to advance the ability to predict ocean dynamics on time and space scales relevant to naval operations. The Department of Energy's (DOE) Office of Energy Research supports marine biological diversity research through its Ocean Margins Program (OMP) and its Microbial Genome Initiative (MGI). The OMP is concerned with assessing the role of the coastal ocean in affecting climate change and the global carbon cycle. In 1994, DOE launched a new molecular biology initiative within the

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--> OMP. The overall goal of this initiative is to provide mechanistic understanding of complex biological processes which mediate major biogeochemical cycles in marine ecosystems, with primary emphasis on the carbon cycle. Molecular biological techniques are being used to address both carbon reduction and carbon oxidation. These molecular biological techniques will be adapted to, and conducted in concert with, OMP field activities scheduled in the southwestern Middle Atlantic Bight. The initiative will emphasize a high degree of coordination and cooperation between investigators, ranging from molecular biologists to "traditional" biologists, chemists, geochemists, and physical oceanographers. The MGI, a spin-off from the DOE/NIH Human Genome Program, will develop a microbial genome sequencing capability that will provide genomic sequence and mapping information on microorganisms with environmental or energy relevance, phylogenetic relevance, or potential commercial importance and application. For the first time, scientists will be able to compare, side by side, genomic sequence information from microorganisms with similar physiological attributes and phylogenetic lineage. This information will further understanding and application of marine biological diversity by providing insight into genetic expression and regulation in key marine microorganisms. Another element in this array of federal activities is the Committee on Environment and Natural Resources (CENR) of the NSTC. The NSTC was established in November 1993 to raise science and technology to the same level of consideration as national security, domestic policy, and the economy. The CENR has been working to develop a national research and development strategy for the federal government on such issues as global change, resource use and management, air quality, toxic substances, natural disasters, and preservation of freshwater and marine environments, as well as to assess relevant social and economic sciences, technologies, and risk assessment techniques. The CENR subcommittees encompass all areas of research on environment and natural resources. One of the CENR's subcommittees, the Subcommittee on Biological Diversity and Ecosystems, is responsible for examining the sustainability of the ecological systems and processes that support life and provide the goods and services necessary for human well-being and opportunity. This subcommittee hopes to develop and promote a research strategy to improve understanding of the interactions among biodiversity, ecosystem dynamics, ecosystem management, and environmental degradation. Finally, the NBS, a newly established component of the Department of the Interior, has a mission of describing the biological resources of the United States (NRC, 1993). One initial charge is to bring all existing inventory knowledge together at a proposed National Biodiversity Information Center. This national agenda for marine biodiversity research can thus work closely with NBS in achieving its goals, and in particular can provide critical marine perspectives as the NBS evolves and grows.