MEANS TO MEASURE BIODIVERSITY
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Conservation Biology and the Preservation of Biodiversity:
The field of conservation biology has been formally recognized for 10, 15, or 20 or more years, depending on how one identifies its beginning (Ehrenfeld 1970; Soulé and Wilcox 1980; Soulé 1986). Regardless of which date we accept, the field has existed for a short time, yet it has had profound and far-reaching effects on the science and management of biodiversity, effects that are well out of proportion to its youthful existence. These influences, some of which I will discuss here, imply that the development of the field of conservation biology was nearly inevitable and perhaps overdue. It brought together and motivated large numbers of scientists of varied description and inclination to address, in a highly pluralistic manner, the problem of the greatest loss of biological diversity in 65 million years. It continues to do so, with some degree of success, although history must be the final judge of its efficacy. I will discuss the field of conservation biology and its contributions to the preservation of biodiversity, identify its areas of weakness, and suggest directions in which the field should go. Much of this material is opinionmy personal assessment of the fieldand little more. It should not be misconstrued as a comprehensive attempt to critically assess the field; that task remains for future analysts.
What is Conservation Biology?
I begin with a general (and admittedly superficial) description of the field; moredetailed treatments are available in many other sources. I offer the definition of conservation biology I have used before (Meffe and Can-oil 1997):
An integrative approach to the protection and management of biodiversity that uses appropriate principles and experiences from basic biological fields such as genetics and ecology; from natural resource management fields, such as fisheries and wildlife; and from social sciences, such as anthropology, sociology, philosophy, and economics.
An important aspect of this definition is that conservation biology borrows and synthesizes from many disciplines. It is an amalgamation of the perspectives, data, techniques, and pursuits of many natural and social sciences, all focused on the problem of the loss and protection of biodiversity
Conservation biology differs from more traditional conservation endeavors, such as fisheries, wildlife biology, forestry, or soil conservation, in at least three ways. First, its origin was strongly academic and theoretical. The field of conservation biology was developed largely by academicians, especially population geneticists and ecologists, who applied their genetic and ecological models to the growing problem of loss of biodiversity. It subsequently was enriched by many other disciplines, in both the natural and social sciences.
Second, the field is rooted in a philosophy of stewardship rather than one of utilitarianism or consumption. The latter has been the basis of traditional resource conservation, that is, conserving resources solely for their economic use and human consumption. This change is reflected in the adoption of very different “guiding lights” in traditional resource management and modern conservation biology: Gifford Pinchot's resource conservation ethic versus Aldo Leopold's evolutionary-ecological land ethic (Callicott 1990).
Third, conservation biology includes significant contributions from nonbiologists in the various social sciences, political sciences, and economics, who join with those in the natural sciences to address our complex problems and develop perspectives and methods. Thus, conservation biology is a broad synthesis of many academic fields, and its purpose is to address the loss and stewardship of biodiversity.
Another important feature of conservation biology is its basis in and recognition of three broad underlying principles (Meffe and Carroll 1997): the inevitability of evolutionary change, the recognition that ecology is dynamic, and the need to take into account the human presence. Conservation biologists recognize that because natural systems are the result of long-term evolutionary change, they will continue to evolve. To protect the status quo, as in a museum, would be a mistake, because systems must continue to evolve. Another mistake is to not understand the evolutionary processes that led to the characteristics of a species when we are attempting to protect or recover it. Likewise, natural systems are dynamic on shorter, ecological time scales, and conservation biologists recognize that natural disturbances are critical to the integrity of ecological systems. The “balance-of-nature” paradigm has been usurped by a “flux-of-nature” viewpoint (Pickett and others 1992).
Finally, conservation biologists recognize that it would be hopelessly naive to ignore humans in the conservation equation or to focus our attention solely on highly natural or pristine systems and lock them away from humanity. In fact, the growing human population is the primary motivation for and the reason we
need the field of conservation biology, and it must be considered at all times. The goal of conservation biology, then, is to understand and meld all three of these foci to help establish an ecologically sustainable world.
What has Conservation Biology Contributed to the Protection of Biodiversity?
I begin this section with a caveat: Although I will claim a great many advances by the field of conservation biology, I do not mean to imply that they all are the result of this field exclusively, that the field has any unique or singular claim to them, or that they would not or could not have developed otherwise. However, I do believe that conservation biology has played an important role in each of these advances.
The major contributions of conservation biology to the protection of biodiversity that I will discuss are of three kinds: new ideas and syntheses, galvanization and reform of natural-resource management, and inspiration for new and related disciplines and current natural-resource practitioners.
New Ideas and Syntheses
First and foremost, conservation biology has provided a formal, global recognition of biodiversitywhat it is and what we are losing (Myers 1992; Wilson and Peter 1988). The world's attention to this crisis and our subsequent modes of dealing with it have been guided largely by this field. In defining biodiversity, we have argued with various degrees of success that biodiversity is much more than richness of species, that it ranges from genes to landscapes and includes the various processes that occur as a function of that diversity. The field has helped to define what we have, what we are losing, and how we deal with it. It is best organized around a triumvirate of composition (what is there), structure (how it is distributed in space and time), and function (what it does) rather than merely around counts of species (Noss 1990). We have learned that if we are to preserve biodiversity successfully, we must deal with natural complexities at multiple levels and configurations.
Next, conservation biology has acted to coalesce many scientific issues under one roof as a metadiscipline. Such issues include genetics, biogeography (including the practical application of island-biogeography theory to rates of loss of biodiversity), population ecology and dynamics, community and ecosystem ecology, evolutionary biology, landscape ecology, and numerous social-science and human dimensions. We consistently draw on these and other disciplines to address the complex interdisciplinary issues that confront us. Conservation biology also recognizes the critical importance of habitat fragmentation and edge effects in losses of biodiversity. It promotes the concept that the quality and spatial configuration of habitat is at least as important to the protection of biodiversity as the total amount of habitat available. Work on metapopulations, spatially explicit models, and the highly practical tool of population viability analysis, also developed by conservation biologists, are related to the spatial considerations of habitat frag-
mentation. All these address the various problems of persistence that face real populations on real landscapes.
Conservation biology has advanced considerably the serious recognition of the potentially disastrous effects of exotic species on native species and ecosystems. The influence of nonindigenous plants and animals has become a major focus in the protection of biodiversity as we have learned how such invaders not only can affect the richness of species, but also can change ecosystem functions.
Finally, conservation biology has incorporated values and ethics into its science. It clearly is a value-laden science,-with a strong value base that freely recognizes that protection of biodiversity is good and necessary, not only for the benefit of humanity, but also for its own inherent good.
Galvanization and Reform of Natural Resource-Management
The second major contribution of conservation biology is that it has acted (whether intentionally or not) to galvanize and reform natural-resource management in two important ways. First, it caused some staid and conservative disciplinessuch as traditional fisheries, wildlife, and forestryto take notice of ideas, controversies, and approaches that had been simmering under the surface for some time. In fact, the clinging to tradition by these fields may have helped to spawn conservation biology, because individuals who were unhappy with the status quo searched for and developed a new discipline that offered an alternative to traditional, consumption-oriented approaches to natural-resource management. As a result, these other disciplines also seem to be moving forward as they embrace the concepts of conservation biology and make them work for natural-resource management. One need only scan the recent pages of such journals as Fisheries and The Wildlife Society Bulletin to see the influence of the last decade of conservation biology.
Conservation biology also has acted in the opposite direction, of bringing ecology and evolution out of the “pure” realm of the ivory tower and applying them to the problems of the day. Many “pure” researchers, who formerly would not dirty their hands with applied problems, now are applying what they know to real landscapes and real issues, thus enriching those endeavors. This new interplay between pure and applied research, with the breakdown of barriers between them, is possibly one of the healthiest and most positive benefits of the development of conservation biology.
Second, conservation biology has changed tangibly management practices as they actually occur on the landscape. In retrospect, the old practices were too scattered and, in many cases, had insufficient scientific justification to have lasted much longer, and their failure may have contributed to the development of conservation biology as a field. As the many pathologies (sensu Holling 1995, Holling and Meffe 1996) of natural-resource management became apparent, new approaches were needed and developed. This has been manifested in several ways:
• challenging and changing natural resource management practices by federal and state agencies to incorporate and accommodate the various principles
promulgated by conservation biology, which now is influencing, through its science and philosophy, how we treat our resources;
• moving away from simple command-and-control approaches to management, which repeatedly have been shown to fail ultimately, toward understanding how nature operates and working within those “rules” (Knight and Meffe 1997);
• developing a greater appreciation for and understanding of uncertainty in environmental management and policy and incorporating that uncertainty into management practices. Recognition of the many natural and human sources of uncertainty has led to multiple calls for adaptive management (Gunderson and others 1995; Holling 1978; Walters 1986), which management agencies are starting to heed and embrace; and
• incorporating natural patterns of variation, such as disturbance regimes, into management. This includes such activities as reinstituting fire in appropriate ecosystems, leaving storm debris on forest floors, and mimicking a natural flood in the Grand Canyon, all designed to incorporate natural processes into management.
In addition to these changes, we are seeing environmental activists working with scientists (or with science) in their calls for policy reform. Numerous activist organizations now routinely incorporate conservation biology into their activities, a step that represents a convergence of science and activism toward the common goal of science-based policy. In sum, natural-resource managers the world over now are relying increasingly on the findings and principles of conservation biology for direction. In the United States, federal and state agencies alike are retooling, using conservation biology as a guide.
Inspiration for New Ideas, Disciplines, and Organizations
New ideas, disciplines, and organizations have been inspired by conservation biology, and a new generation of practitioners is undergoing intellectual development and professional training in this new environment. For example, the idea that cross-boundary issues are critical is now a common point of discussion among natural-resource management agencies and private landowners, whereas 10 years ago, political boundaries on a map seemed real and impermeable. Stepping back to view the larger landscape and cooperate with other land tenants, rather than hiding behind the seemingly comfortable and protective boundaries set by legal documents, is becoming a way of life rather than an unusual behavior. In general, such notions of cooperation for a common good rather than of confrontation or competition, are becoming prevalent.
New disciplines have been denned or developed further as a result of progress within conservation biology. For example, restoration ecology, landscape ecology, environmental ethics, and ecological economics all have begun to flourish as important components of conservation biology. Surely they existed beforehand, and they may have developed independently, but conservation biology seems to have been and continues to be the overarching catalyst that supported and promoted
their advancement. The metadiscipline of conservation biology is the glue that binds these and other disciplines into a coherent and focused package.
An obvious but extraordinarily important catalyst for the field was development of a major international society, the Society for Conservation Biology, and its journal, Conservation Biology, as the focal points for intellectual activities in the field. The effects and influences of this society and journal are virtually incalculable within the academic and applied communities of conservation scientists. They help to identify and define the field and offer an intellectual home to its practitioners.
Closely related to all these factors, and ultimately feeding the further development of conservation biology, are the many courses and degree programs in conservation biology that are developing in colleges and universities around North America and the rest of the world, as well as several college textbooks that are designed specifically for use in these courses. For the first time in history, the field has reached the point at which we are formally educating a new generation of students as conservation biologists, in contrast with the founding generation, who came to the field from various specialized disciplines. These students have been inspired by the challenges and opportunities involved in the protection of biodiversity, which seems to have given greater meaning to basic programs in ecology.
Finally, training courses have developed in various natural-resource management agencies to bring the practitioners up to speed on such topics as general tenets of conservation biology, ecosystem management, and various human dimensions. My experiences as a trainer in some of these courses tells me that as a result of this reorientation natural-resource management in the United States will never be the same.
Gaps and Problems
Although conservation biology has been considered a rousing success by most measures, it has its problems, it has experienced growing pains, and it still has some way to go to be considered a mature discipline. A useful analogy is human ontogeny. Conservation biology was born rapidly, with typical pains and shakiness; it grew quickly, feeling its way along, learning first how to walk, then to run; it became an awkward adolescent; and now it is emerging into confident maturity as a young adult. It has not reached its full potential yet, and it has a great deal to learn before it has its full effect on the world, but its future looks bright and exciting. However, hurdles must be overcome, and I present several of them here.
First, I believe the field's main problem is that it means very little globally, compared with many other human endeavors; conservation biology certainly is not yet a household term that most people can identify. Society at large does not realize what conservation biologists have to offer or the relevance of conservation biology to their lives, other than in a vague connection to a general concern for the environment. Conservation biologists have not done a good job of positioning the field to be a globally effective agent of social change.
Part of the problem is that society has not defined its environmental problems broadly enough to address them adequately. Many individuals seem to associate environmental problems with the need to recycle, with possible global climatic change, with harm to individual animals, with the problem of toxins in the air, water, and soil, or with other issues related to human health. As important as these problems are, otherssuch as major losses of biodiversity and their ramifications, collapse of ecosystem services, and destruction and fragmentation of habitat (apart from tropical deforestation, which much of the public recognizes)do not seem to resonate as major environmental issues or issues that hold much threat for or relevance to humanity. Many people do not seem to make connections between the development of strip malls or golf courses, growth of population, loss of soils, withdrawal of water, and related activities and their influences on biodiversity, sustainability, human health, or social structure. In essence, I do not believe that society at large appreciates what really supports human populations or why desertification, logging of old-growth forests, and mass extinction of species are critically important to all peoples.
Second, the field of conservation biology developed with a largely terrestrial bias, which it retains. Consequently, it has lagged in addressing problems in freshwater aquatic systems and, especially, marine systems (Irish and Norse 1996). Recent attention to the marine realm, including major marine symposia at the meeting of the Society for Conservation Biology in 1997, seems to be addressing that problem.
Third, in my opinion, conservation biology is still too academic: it clings to its roots in academe and seems fearful of venturing too far into unknown territories. I believe that conservation biologists need to be more pragmatic and more practical, and the field needs more relevance to immediate problems of the day if we are to have a greater influence on the protection and recovery of biodiversity. To do this, we must dare to leave the comfort of the academic womb and take greater risks in the real world of conservation action.
Fourth, the nature of the university system itself, at least in the United States, has done little to foster risk-taking and creativity and much to promote conservatism and the status quo. With its high disciplinary walls (Meffe 1998), adherence to tradition, and rewards for conformity, academe not only frustrates progress in a new discipline, such as conservation biology, but does little to address the major environmental and social problems of the day (Orr 1994). Rather than playing a leadership role in cutting-edge ideas, universities often seem to lag behind, restricting such activities and rewarding those which bring in large sums of money for low-risk work. Much of the activity in conservation biology is taking place outside universities, in resource-management agencies, advocacy groups, and even resource-extraction industries.
Where does the Field Need to Go?
I think the field should move in several directions and be strengthened in some areas so that conservation biology can develop further as a discipline and, more importantly, be able to influence society with more scientifically based decisionmaking.
• Conservation biology needs greater synthesis with other disciplines or subdisciplinessuch as restoration ecology, design (broadly denned to include all human-made products and endeavors), and ecological economicsand with various human dimensions such as sociology, psychology, and anthropology. Conservation biology has something to contribute to all these, and vice versa. Greater communication across fields, some where conversations possibly have never occurred, can help to promote problem-solving in the broadest sense.
• Conservation biologists need to do a better job of teaching about the connection between the overall ecological condition and individual human-health or social conditions. Many times, the arguments we muster for the protection of biodiversity, although compelling to scientists, do not resonate with average citizens who are just trying to make a living. In addition to the various moral arguments we typically use to justify our concerns, we need to do a better job in making it clear that functional natural ecosystems are necessary for workable human social systems and the health and vigor of all humankind. Conservation biology is concerned not just with nature, but very much with humanity as well.
• We also need to take the lead in modifying educational curriculafrom kindergarten through graduate levelto reflect better the central importance of an ecological perspective in society. The primary task will be to break down the artificial disciplinary boundaries that have haunted education for centuries, to overcome departmental territorialities, and to cease the extreme specialization that so often results in narrow technical training rather than a broad education that can lead one to understand the interrelationships in complex problems and begin to address them. We need to stop teaching as though mathemathics, sociology, biology, engineering, history, and literature are unrelated We need to do a better job of teaching the full diversity of the human experience and of centering it on functioning ecosystems that make the planet livable for all species, including humans.
• Conservation biology has a golden opportunity to join with many and varied religious interests that focus on environmental awareness and protection of life on Earth. For example, so-called “green evangelicals” fervently recognize and understand the importance of protecting biodiversity, although the term they use is different (“God's creations”). Seeing all life as the result of a single event of creation and interpreting Biblical writings on dominion as a responsibility for stewardship rather than a license for domination and control of nature, this perspective can be valuable beyond measure, reaching large numbers of people who otherwise might not identify with “biodiversity” or care much about it from a scientific perspective. Harnessing the energy of religious perspectives concerned with guardianship of creation can be a powerful boost to protection of biodiversity.
• Most important, I think, we need to do a better job of incorporating what we know into effective public policy. We need to make our science work; we need to put it to daily use. It is time for conservation biology to move to a new plateau in society, to make our presence known, our science relevant, and our views sought and respected. Ideally, the public should listen to what conservation biologists have to say with as much anticipation, concern, and enthusiasm as they have for daily stock-market reports, economic forecasts, or news about medical advances.
I believe the science of conservation biology is in an extremely active, turbulent, and exciting period of development right now. The last 15 years have seen dramatic changes in conservation priorities, techniques, philosophies, and approaches. Now is when the science is being molded, when the approaches to the enormous challenges to humanity are being mapped out, and when the future of biodiversity and humanity largely are being determined. This is a thrilling, frightening, and wonderful time to be practicing conservation science, one that I hope we can look back on with pride and satisfaction. Conservation biology has taken huge strides in the effort to protect biodiversity, but these still are only the initial, cautious steps of a long and never-ending journey; we have much yet to learn and accomplish.
Callicott JB. 1990. Whither conservation ethics? Cons Biol 4:15–20.
Ehrenfeld DW. 1970. Biological conservation. New York NY: Holt, Rinehart and Winston.
Gunderson LH, Holling CS, Light SS (eds). 1995. Barriers and bridges to the renewal of ecosystems and institutions. New York NY:Columbia Univ Pr.
Holling CS. 1978. Adaptive environmental assessment and management. New York NY: J Wiley.
Holling CS. 1995. What barriers? What bridges? In: Gunderson LH, Holling CS, Light SS (eds). Barriers and bridges to the renewal of ecosystems and institutions. New York NY: Columbia Univ Pr. p 3–34
Holling CS, Meffe GK. 1996. Command and control and the pathology of natural resource management. Cons Biol 10:328–37.
Irish KE, Norse EA. 1996. Scant emphasis on marine biodiversity. Cons Biol 10:680.
Knight RL, Meffe GK. 1997. Ecosystem management: agency liberation from command and control. Wildl Soc Bull 25:676–8.
Meffe GK. 1998. Softening the boundaries (editorial). Cons Biol 12:259–60.
Meffe GK, Carroll CR. 1997. Principles of conservation biology, second edition. Sunderland MA: Sinauer.
Myers N. 1992. The primary source. New York NY: WW Norton.
Noss RF. 1990. Indicators for monitoring biodiversity: a hierarchical approach. Cons Biol 4:355–64.
Orr DW. 1994. Earth in mind: on education, environment, and the human prospect. Washington DC: Island Pr.
Pickett STA, Parker VT, Fiedler PL. 1992. The new paradigm in ecology: implications for conservation biology above the species level. In: Fiedler PL, Jain SK (eds). Conservation biology: the theory and practice of nature, conservation, preservation, and management. New York NY: Chapman & Hall. p 65–88.
Soulé ME, Wilcox BA. 1980. Conservation biology: an evolutionary-ecological approach. Sunderland MA: Sinauer.
Soulé ME (ed). 1986. Conservation biology: the science of scarcity and diversity. Sunderland MA: Sinauer.
Walters CJ. 1986. Adaptive management of renewable resources. New York NY: McGraw-Hill.
Wilson EO, Peter FM (eds). 1988. Biodiversity. Washington DC: National Acad Pr.
Applying Molecular Methods to Maximize the Conservation of Taxonomic and Genetic Diversity.
Conservation biology is an applied science that involves direct human intervention into the management of diminishing natural resources. However, unlike traditional resource management, the focus of conservation biology is not necessarily driven by direct economic incentive or the desire to manage a resource for the sake of harvesting it. Instead, the primary goal of this field is to stop the downward spiral of loss of biological diversity by mitigating factors that erode the biological integrity of intact ecosystems and the long-term evolutionary viability of populations, species, and communities of organisms. In this sense, conservation biologists attempt to manage biodiversity on two time scales, the ecological (present) and the geological (future), but management itself is inevitable.
By definition, conservation biology must be multidisciplinary, requiring an integration of many areas of biology, including biogeography, systematics, plant and animal ecology, reproductive biology and physiology, range and wildlife management, environmental toxicology, population biology, genetics, and molecular biology. Moreover, the coordination or planning of any conservation effort also involves issues outside the realm of biology, because most environmental-conservation solutions are compromises between the biological requirements of a natural system and the socioeconomic and political realities of the human populations that are associated with that system. Therefore, the conservation of biodiversity must strike a balance between the needs of a growing human population and the viability of biological systems in the face of a rapidly changing environment.
The act of preserving our natural or biological resources, as in so many action-oriented fields, can be distilled into five simple questions: why, what, where, how,
and who? These questions are simple, but their answers have proved vexing and indeed have generated some intense debates. The answer to the first and perhaps most important societal, issue, why, has been framed in ways that range from the economics of ecosystem services to the psychological and cultural value of intact ecosystems and species to our moral and ethical obligations to pass on to our children the natural world in roughly the same condition in which we found it (Costanza and others 1997; Kellert and Wilson 1993; Pimm 1997). A detailed discussion of these issues is outside the scope of this paper; we will assume that the reader will find justification elsewhere for why we should conserve our natural resources.
Assuming we should develop a rational means to describe and conserve the world's biological diversity, we must rely on some scientific systems of measurement and theory to address the four remaining critical questions. Over the last 10 years, conservation genetics has emerged as a subfield of biological conservation that offers an objective approach to three of these questions: what, where, and how. Conservation genetics is more a focus than a field of study, but it has at its root the application of molecular and quantitative genetic methods to the preservation of genetic, species, and ecosystem diversity. Genetics can be applied to these issues in numerous ways, but the term conservation genetics usually refers specifically to molecular genetic techniques that help to
• identify evolutionarily distinct groups of organisms (for example, populations or species) that are worthy of separate conservation efforts (that is, conservation units) (Avise 1996; Moritz 1994, 1995; O'Brien 1996);
• define specific geographic regions that harbor genetically distinct populations, and/or species (that is, regions of genetic endemism) (Avise 1996; Riseberg and Swensen 1996; Templeton and Georgladis 1996; Vane-Wright and others 1994; Williams and Humphries 1994; Witting and others 1994); and
• estimate the distribution of genetic diversity within and among conservation and management units to develop plans that will conserve the greatest amount of that diversity and the evolutionary potential it offers (Burgman and others 1993; Caughley 1994; Frankham 1995; O'Brien 1994).
Researchers have used genetic approaches to address a variety of conservation problems in plants and animals found on many continents. This genetic research has involved diverse laboratory procedures and approaches to data analysis; the results have provided critical information for wildlife managers and environmental policy-makers. In the following sections, we present a brief overview of both the methods used in conservation genetics and some empirical studies that underscore the value of genetic analysis in conservation management.
what are the Conservation Units?
A critical first step in designing appropriate conservation measures is properly defining and identifying the group one wishes to conserve, the so-called conser-
vation unit (CU). In other words, we need to define what we wish to conserve before we can take measures to conserve it. Although this may seem trivial, the evolutionary process is often murky enough to lead to great difficulty in defining CUs, particularly when we are dealing with closely related species that have been thrown together recently by human-induced changes in the environment or those which were never isolated fully from one another reproductively (that is, hybridization has occurred). In these cases, careful examination of the biological characteristics of an organism that are most likely to carry evolutionary historical information (a field known as biological systematics)such as features of anatomy, behavior, and geneticsoften yields the clues necessary to place a series of populations and species on a synthetic family tree (also known as a phylogenetic tree). These clues even can be used to determine whether a group of organisms can be defined as a single evolutionarily significant unit (ESU), which may be but is not necessarily synonymous with what we usually refer to as a species.
Molecular-generic approaches to biological systematics have emerged as one of the most exciting new areas of biological research (Hillis and others 1996). A wide array of technical and analytical methods has been used to address issues of evolution and conservation at all levels of organization, ranging from genes within populations through the process of speciation to the reconstruction of the tree of life itself (Avise 1994). Of particular importance here are the contributions of molecular techniques to the identification and phylogenetic placement of rare and endangered species. Knowledge about diversity at the molecular level can be used to reconstruct the evolutionary history of an endangered organism (Avise and Hamrick 1996) and to identify the ESUs on which to focus our attention for conservation (Moritz 1994, 1995). Because much of conservation planning depends on taxonomic or species assignments (Taberlet 1996), identifying systematically based CUs aids considerably in developing management plans and in evaluating priorities for conservation (Smith and Wayne 1996).
Molecular-systematic studies can help clarify taxonomic issues at three different levels. First, we can identify cases of “oversplitting”, that is, when distinct morphological forms are considered different evolutionary entities but are in fact genetically indistinguishable. This implies that gene flow still may be occurring between the different forms and that they therefore should not be considered evolutionarily distinct. For example, the now-extinct dusky seaside sparrow (Ammodramus maritimus nigrescens) of Florida, originally described as a distinct species, was redefined later as a subspecies when it was shown to be genetically indistinguishable from other populations of seaside sparrows (Avise and Nelson 1989). Moreover, it was shown that all populations of seaside sparrows on the Atlantic coast (including the dusky seaside sparrow) are genetically more similar to each other than they are to populations of seaside sparrows that are found along the Gulf of Mexico. Clearly, molecular data supported the inclusion of the dusky seaside sparrow in the seaside-sparrow species and suggested that its loss, although regrettable, had little or no effect on the long-term evolutionary course of the entire species.
Second, molecular systematics can help us distinguish between forms that are morphologically similar but are in fact ancient, unrelated lineages with little or no gene flow between them. One example is Darwin's fox (Dusicyon fulvipes) on Chiloé Island in Chile. Some scientists had considered Darwin's fox a small race of the common South American grey-fox species Dusicyon griseus, on the basis of morphological similarities. Chiloé island is only 5 km off the coast and likely was connected to the mainland during the last glaciation (about 13,000 years ago), which would have created opportunities for gene flow between Darwin's and grey foxes. However, genetic analyses of Darwin's fox and other South American fox species suggest that Darwin's fox is at least as divergent from the grey fox as the grey fox is from another well-recognized fox species, the culpeo fox (Dusicyon culpaeus), and that Darwin's fox probably evolved from the first immigrant foxes into South America 2–3 million years ago. Recently, a small population of Darwin's foxes was found on mainland Chile, and they were shown to be quite genetically divergent from the grey fox but closely related to the population on Chiloé island. This suggests little or no present or historical gene flow between Darwin's and grey foxes, and it supports the distinctiveness of Darwin's fox as a separate species (Wayne 1996).
Third, systematic analyses of genetic characters can provide an objective means of identifying evolutionarily distinct lineages among closely related groups. The Iberian lynx, Lynx pardinus, is considered to be the most vulnerable cat in the world. Its remaining populations are highly fragmented and of limited size. The species status of the Iberian lynx is complicated: Some consider it to be a geographic variant of the Eurasian lynx, Lynx lynx, and others consider it to be a distinct species. Because the taxonomic status of the Iberian lynx is important to the establishment of an effective management plan for lynxes in general, a molecular-systematic study was conducted recently (Beltran and others 1996). The results of this study revealed a close relationship between the Canadian lynx (Lynx canadensis) and the Eurasian lynx, but the Iberian lynx is evolutionarily more distinct. Thus, these molecular data give validity to the concept that the Iberian lynx is a phylogetically distinct species that deserves separate consideration for conservation.
Where do those Conservation Units Reside?
Once we decide what groupings of organisms are distinct and worthy of separate efforts at conservation (that is, we identify our CUs), it becomes critical to determine the geographic location of important subsets of individuals within each CU. In other words, where will we focus our conservation efforts to preserve a CU or species?
The use of molecular systematics in a geographic context can contribute to answers to this question in two ways. First, detailed studies of intraspecific (within-species) variation can identify the geographic limits of either a CU or what Moritz (1994) calls a management unit (MU). Second, patterns of intraspecific phylogenies of unrelated groups of organisms may assist in identifying geographic regions
whose populations and species have had a shared, unique evolutionary history, thus allowing for the conservation of communities of organisms that have high levels of genetic endemism or uniqueness.
The term intraspecific phylogeography denotes the connection between biological systematics, population genetics, and biogeography, the study of the distribution of organisms in geographic space and the factors that led to that distribution (Avise and others 1987). In principle, any biological characteristic can be used for this purpose, but intraspecific phylogeography now mostly is associated with the study of molecular markers, especially mitochondrial DNA (in animals) or chloroplast DNA (in plants). By determining the detailed genetic and evolutionary relationships of populations within a species (or CU), and superimposing that intraspecific molecular phylogeny on a geographic map, one can infer the processes that historically determined the current distribution of organisms. One also can use this approach to identify the geographic location of genetically distinct populations (that is, populations that substantially differ from one another by the frequency of genetic traits rather than by the presence or absence of those traits) or MUs, which might deserve special attention if specific conservation measures become necessary to preserve a given species. The identification of specific MUs and their geographic location currently has one of the highest priorities in most efforts that use molecular markers for conservation purposes, and intraspecific phylogeography provides a theoretical framework to accomplish this.
Case Study. Among the animal species currently listed by the International Union for the Conservation of Nature (IUCN), the Convention on International Trade in Endangered Species (CITES), and the US Department of the Interior as endangered, the Sumatran rhinoceros (Dicerorhinus sumatrensis) is one in greatest need of special attention and immediate wild-population (or in situ) management. Historically, this species inhabited most of the Indochinese peninsula, from Burma (Myanmar) to Vietnam, and south to Malaysia and the islands of Sumatra and Borneo. Destruction of habitat and hunting have led to a rapid decline of this species over the last 2 decades. Only a few confirmed populations remain on peninsular Malaysia, Borneo, and Sumatra. Because of the dire situation of this species, translocation programs have been proposed that would move individuals that are scattered among fragments of unsustainable forest and concentrate them in protected zones of natural habitat (Foose and van Strien 1995). However, it is important to remember that the objectives of any conservation effort should be not only to maintain a collection of organisms, but also to preserve the maximal amount of existing genetic variability within a species and to maintain the evolutionary historical integrity of its wild populations.
Geographic mapping of the distribution of mitochondrial-DNA (mtDNA) variants among Sumatran rhinoceros populations (Morales and others 1997), using both molecular-systematic and population-genetic methods, reveals two phylo-geographic features that are important to the conservation of the Sumatran rhinoceros. First, a phylogenetic tree of mtDNA haplotypes, overlaid on the distri-
bution of the Sumatran rhinoceros, suggests that the population in Borneo possesses a unique mtDNA variant that is not shared with other Sumatran rhinoceroses, indicating a long history of isolation from the remaining Sumatran and peninsular Malaysian populations. Therefore, the population in Borneo should be considered a separate ESU. Second, a geographically referenced population-genetic analysis suggests that the populations outside Borneo can be divided into two groupings or MUswest Sumatra, and east Sumatra and Malayaon the basis of significant differences in the frequency of mtDNA variants and the restriction of gene flow that they imply. Thus, translocation and other conservation efforts should take these three distinct units (ESUs or MUs) into consideration and try to maintain the evolutionary and genetic integrity of each unit.
More recently, and closely related to the common use of intraspecific phylogeography, efforts have been made to map intraspecific phylogeographic patterns simultaneously among a wide variety of species that occupy overlapping geographic ranges. This has been done to identify regions that harbor populations or species that are consistently genetically distinct from other populations within their species (so-called conspecifics) or other closely related (sister) species within their genus (so-called congenerics). These regions of genetic uniqueness or genetic endemism can be used to design reserves and other mechanisms of conservation (Avise and Hamrick 1996; Templeton and Georgladis 1996; Williams and Humphries 1994; Witting and others 1994). They also provide an effective shortcut to making decisions about conservation at the levels of species and community because it would be impossible to conduct individual genetic surveys of the hundreds of thousands of species in a particular region. Thus, a consistent pattern of regional genetic uniqueness across a diverse but logistically feasible number of species (including fungi, plants, invertebrates, and vertebrates) would allow one to assume reasonably that most populations or species within that particular region, the large majority of which will not have been analyzed, are genetically unique.
Case Studies. One compilation of several studies of invertebrate and vertebrate animals of the southeastern United States found major patterns of molecular phylogeographic congruence among populations of coastally distributed species (Avise 1996). These patterns were shared among varied groups of organisms, including horseshoe crabs, American oysters, diamondback terrapins, ribbed mussels, seaside sparrows, toadfish, black sea bass, and tiger beetles. Even species that have a greater ability to disperse, like white-tailed deer, showed a similar pattern, suggesting population differentiation in this region in response to a persistent set of historical biogeographic processes (Ellsworth and others 1994). The pattern revealed by most species indicates major molecular phylogenetic discontinuities between the Gulf of Mexico and the Atlantic coastline of the southeastern United States, whereas some patterns, like that seen for deer, indicate the uniqueness of the populations in southern Florida. Together, these findings suggest that maritime and other species in this region may have been subjected to the same bio-
geographic influences and thus share a common biogeographic history. Although some exceptions to the common molecular-phylogeographic pattern exist, the evidence is strong that at least among animal species, populations on either side of southern Florida are likely to be genetically distinct from one another. The conservation of any one of those species must incorporate efforts on both sides of this important biogeographic divide.
How will We Manage Genetic Diversity within each Conservation Unit?
Once we have determined what groupings of organisms (CUs) are distinct and worthy of separate efforts at conservation, and where the genetically unique populations of those units (MUs) or larger regions of general genetic uniqueness (regions of genetic endemism) are, it is critical for us to devise the means to conserve those species individually or regionally. In other words, how will we conserve each species's populations and their underlying genetic diversity well into the future?
Population genetics offers a key perspective on this issue because most critical evolutionary events occur at the level of the population. The potential rate of evolution depends on the amount of genetic diversity in a population; processes that erode levels of genetic diversity or increase the occurrence of deleterious combinations of genes (such as inbreeding) within populations limit the rate and scope of potential evolutionary changes in those populations to meet environmental challenges (Templeton and others 1990). Furthermore, biologists agree that levels of genetic diversity within individuals may confer important advantages of fitness on those individuals (Allendorf and Leary 1986). Thus, a fundamental concern of conservation biologists is to preserve genetic diversity in populations and species and the resulting evolutionary potential. The field of population genetics plays a critical role in determining how that diversity is distributed and how best to preserve it.
Population Genetic Structure
A species's genetic diversity can be distributed in various ways, depending on historical ecological, geological, and human-induced events, as well as on the current patterns of geographic distribution, individual dispersal, social organization, ecological adaptation, demographic transition (births, deaths, and generation length and overlap), and genetic migration (the flow of genes across a landscape). How one configures a conservation-management strategy to encompass the individuals and populations that are necessary to capture the greatest amount of a species's genetic diversity will be derived largely from knowledge of the existing distribution of that diversity across the species's range, otherwise known as the population genetic structure of a species. Information of this sort is essential for conservation planning but is often difficult to obtain.
Case Study. Among Asian macaque monkeys are species that have extensive geographic distributions, either contiguous through the mainland, like the rhesus
monkey (Macaca mulatto), or fragmented, like the long-tailed macaque (Macaca fascicularis), whose distribution includes part of mainland Asia and many islands in the Malay Archipelago and the Philippines. Melnick (1988) and Melnick and Hoelzer (1992) have shown that 91% of the nuclear genetic variation in the rhesus monkey can be attributed to variation within a geographic region; in the more fragmented long-tailed macaque, this figure is reduced to 67%. In other words, nearly 4 times as much of a species's genetic variation can be attributed to differences between regions in the more fragmented long-tailed macaque as in the more contiguously distributed rhesus monkey. This pattern also holds true for species that have a much more restricted distribution, like the toque macaque (Macaca sinica) of Sri Lanka and the Japanese macaque (Macaca fuscata). In the toque macaque, which exists on only one island, only 3% of the species variation can be attributed to differences between regions, whereas in the Japanese macaque, which exists on a number of islands in Japan, that figure increases to 24%.
What does all this mean in terms of conservation? Very simply, the greater the percentage of overall genetic variation in a species that can be attributed to differences between populations, the greater the number of populations that must be included in a conservation-management plan that seeks to preserve some maximal level (for example, 90%) of existing genetic diversity. The Japanese macaque is considered an endangered species by the IUCN; thus, given its current population genetic structure, efforts to conserve this species must include a broad geographic representation of different island populations to maximize the genetic diversity to preserve. If the toque macaque ever shares a similar fate in Sri Lanka, reserves that harbor only a small number of sufficiently large populations likely will capture most of the species's existing variation.
As human populations continue to grow, landscapes that are fragmented by human activities are becoming the predominant arena within which demographic and evolutionary processes in terrestrial plants and animals occur. Nevertheless, the effects of human-induced changes in the landscape on the distribution of genetic variation in wild populations remain largely unknown. It is important to examine the long-term genetic consequences of fragmentation of habitat so we can develop appropriate strategies for maintaining viable populations in remnants of habitat over hundreds to thousands of years. Such studies are only beginning to be conducted, but the new area of metapopulation analysis and management has emerged as a result of these issues (Hanski and Gilpin 1997). A metapopulation is characterized as a network of populations that have limited gene flow between them and have population extinction and recolonization in specific localities (Levins 1969). In the context of management, we define a situation as extinction if a population either has died out or has been removed for the purpose of translocation.
Metapopulation management brings together the fields of demography, population genetics, and resource management. The primary goal is to “fool” the evolutionary process into “believing” it is acting on one large contiguous population, with all the attendant complexities of births, deaths, dispersal, and local group ex-
tinctions, when in fact the members of the species are distributed in many small patches that effectively are isolated from one another by intervening, unsuitable habitat through which they cannot cross. In general, a-metapopulation-management plan involves in-depth study of the behavior, demography, and genetics of a species to determine when, how many, and where individuals should be moved among the existing patches of suitable habitat so as to mimic one large panmictic (free-mixing) population or species. The critical long-term goal of such a strategy is to provide the largest number of breeding individuals, or effective population size, to maintain most of the population's or species's genetic diversity over the course of centuries (Wade and McCauley 1988). One can demonstrate both mathematically and experimentally that the larger the effective population size, the less likely that genetic variation will be lost to random processes that generally remove genetic variants from a population (so-called genetic drift). Hence, the general goal is to maintain as large an effective population as possible, thus buffering the forces that otherwise would inevitably erode genetic diversity.
Case Studies. Populations of the ocelot (Leopardus pardalis) in southern Texas provide an example of a habitat specialist that has been fragmented into many small populations after 50 years of converting land to agricultural uses. A recent examination of genetic variation has revealed a lack of gene flow between the populations in southern Texas and the historical source population in northern Mexico (Walker 1997). Furthermore, genetic variation within populations in southern Texas has eroded. Assuming that a generation in the ocelot is about 2 years, this means that the fragmentation of the ocelots' range into small, relatively isolated populations has resulted in a major loss of genetic variation in only 25 generations. If genetic diversity within populations of ocelots in southern Texas is to be restored and maintained, any future conservation plan must involve exchanging cats between these isolated populations and those in northern Mexico.
Black lion tamarins (Leontopithecus chrysopygus), endemic to the state of São Paulo, Brazil, exist in only seven forest fragments (Coimbra-Filho 1976; Valladares-Padua 1993). Researchers from the Instituto Pesquisas Ecolõgicas (IPÊ) in São Paulo and from the Center for Environmental Research and Conservation (CERC) at Columbia University in New York are undertaking a project to devise a program of metapopulation (translocation) and management for these animals. The immediate goal of this effort is to translocate individuals from one forest fragment to another. The ultimate goals are to ensure proper assimilation of introduced individuals into other populations or into unoccupied but suitable patches of habitat and to conserve a “natural” amount of genetic diversity in the combined forest fragments, including the empty ones that will be colonized by translocated lion tamarins. One way to ensure proper assimilation of introduced lion tamarins is to mimic their natural dispersal patterns and their current population genetic structure. The genetic data from this study will contribute immeasurably to what is known about the social organization, dispersal patterns, and population genetics of the black lion tamarin and thus will enhance the chances of successful translocation, demographic stability, genetic management, and long-term survival of this highly endangered species.
Conservation Genetics Training
Who will Perform Genetic Analysis for Conservation?
Conservation-management recommendations that come from outside the nation in which they are to be implemented rarely are followed. Indeed, innumerable counterexamples teach us that conservation is done best when it is done at home. For this reason, scientists from each country in which the work is to be done should be trained as conservation geneticists. Thus, when asking who should be performing genetic analysis for conservation, the logical answer would be the countries' scientists who would use the resulting genetic information to establish and revise their conservation programs and policies.
This means that, in addition to training our own students and future conservation geneticists, the universities and other research institutions in developed countries should be providing opportunities for in-depth technical and analytical training to young scientists who have the best chance of establishing this type of research in their own countries. One program that is doing just that is the Conservation Genetics Training Program for Southeast-Asian Scientists, which is based at CERC (see Melnick and Pearl 2000) and is funded by the MacArthur Foundation. This year-long program provides training in research design, laboratory techniques, and data analysis. As a followup to this training, CERC staff help the trainees establish research programs in their home countries. This assistance ranges from technical guidance to the actual purchase and outfitting of small laboratories to do the work. This program has trained researchers from Indonesia, Malaysia, Thailand, Vietnam, and China, and the CERC training staff now includes a postdoctoral scientist from Sri Lanka. Out of this program is a rapidly growing regional cadre of researchers who publish in peer-reviewed international journals (Wang and others 1997). This group is likely to have a major effect on future decisions about conservation within Southeast Asia.
In this chapter, we have highlighted the important uses of genetic analyses to define the units of conservation and the units of management, the geographic locations of those units, and the ways in which genetic variation is distributed within and among the populations that make up each unit. This discussion and the examples we have offered are meant to provide a brief introduction to the nonspecialist reader and to highlight the value of these approaches for wildlife managers, other conservation practitioners, and environmental policy-makers. It is important, however, to point out that other biological disciplines (for example, morphological systematics and behavioral ecology) also contribute significantly to the definition of evolutionary distinctiveness and that many other considerationssuch as overall evolutionary uniqueness, current vulnerability, and socio-cultural valuemust be considered when we are developing protective measures for a particular population or species. Ultimately, we must apply as much information as possible to decisions about designing and launching conservation efforts. We hope it is clear that genetic analysis is a powerful and timely mechanism for generating a great deal of valuable information for the purpose of conservation.
Allendorf FW, Leary RF. 1986. Heterozygosity and fitness in natural populations of animals. In: Soulé ME (ed). Conservation biology: the science of scarcity and diversity. Sunderland MA: Sinauer. p 57–76.
Avise JC, Hamrick JL (eds). 1996. Conservation genetics: case histories from nature. New York NY: Chapman & Hall. 512 p.
Avise JC, Arnold J, Ball RM, Bermingham E, Lamb T, Neigel JE, Reeb CA, Saunders NC. 1987. Intraspecific phylogeography: the mitochondrial DNA bridge between population genetics and system-atics. Ann Rev Ecol Syst 18:489–522.
Avise JC, Nelson WS. 1989. Molecular genetic relationships of the extinct dusky seaside sparrow. Science 243:646–8.
Avise JC. 1994. Molecular markers, natural history, and evolution. New York NY: Chapman & Hall. 511 p.
Avise JC. 1996. Toward a regional conservation genetics perspective: phylogeography of faunas in the southeastern United States. In: Avise JC, Hamrick JL (eds). Conservation genetics: case histories from nature. New York NY: Chapman & Hall. p 431–70.
Beltran JF, Rice JE, Honeycutt RL. 1996. Taxonomy of the Iberian lynx. Nature 379:407–8.
Burgman MA, Ferson S, Akqakaya HR. 1993. Risk assessment in conservation biology. New York NY: Chapman & Hall. 314 p.
Caughley G. 1994. Directions in conservation biology. J Anim Ecol 63:215–4.
Coimbra-Filho AF. 1976. Os saguis do genero Leontopithecus Lesson, 1840 (Callithicidae-Primates).
Unpublished Master's thesis, Universidade Federal do Rio de Janeiro.
Costanza R, d'Arge R, de Groot R, Farber S, Grasso M, Hannon B, Limburg K, Naeem S, O'Neill RV, Paruelo J and others. 1997. The value of the world's ecosystem services and natural capital. Nature 387:253–60.
Ellsworth DL, Honeycutt RL, Silvy NL, Bickham JW, Klimstra WD. 1994. Historical biogeography and contemporary patterns of mitochondrial DNA variation in white-tailed deer from the southeastern United States. Evolution 48:122–36.
Foose TJ, van Strien NJ (eds). 1995. Asian rhinos. Newsl IUCN SSC Asian Rhino Specialist Group: No 1.
Frankham R. 1995. Conservation genetics. Ann Rev Gen 29:305–27.
Hanski IA, Gilpin ME (eds). 1997. Metapopulation biology, ecology, genetics, and evolution. San Diego CA: Academic Pr. 512 p.
Hillis DM, Moritz C, Mable BK. 1996. Molecular systematics, 2nd ed. Sunderland MA: Sinauer. 655 p.
Levins R. 1969. Some demographic and genetic consequences of environmental heterogeneity for biological control. Bull Entomol Soc Amer 15:421–31.
Kellert SR, Wilson EO (eds). 1993. The biophilia hypothesis. Washington DC: Island Pr. 439 p.
Melnick DJ. 1988. The genetic structure of a primate species: Rhesus macaques and other cercopithecine monkeys. Inti J Primat 9:195–231.
Melnick DJ, Hoelzer GA. 1992. Differences in male and female macaque dispersal lead to contrasting distributions of nuclear and mitochondrial DNA variation. Intl J Primat 13:379–93.
Melnick DJ, Pearl MC. 2000. Center for Environmental Research and Conservation (CERC): a new multi-institutional partnership to prepare the next generation of environmental leaders. In: Raven PH, Williams T (eds). Nature and human society: the quest for a sustainable world. Washington, DC: National Academy Press, p. 462–70.
Morales JC, Andau PM, Supriatna J, Zainuddin ZZ, Melnick DJ. 1997. Mitochondrial DNA variability and conservation genetics of the Sumatran rhinoceros. Cons Biol 11:539–43.
Moritz C. 1994. Defining evolutionarily significant units for conservation. Trends Ecol Evol 9:373–5.
Moritz C. 1995. Uses of molecular phylogenies for conservation. Phil Trans R Soc London Series B 249:113–8.
O'Brien S. 1994. A role for molecular genetics in biological conservation. Proc Natl Acad Sci USA 91:5748–55.
O'Brien S. 1996. Conservation genetics of the Felidae. In: Avise JC, Hamrick JL (eds). Conservation genetics: case histories from nature. New York NY: Chapman & Hall. p 50–74.
Pimm SL. 1997. The value of everything. Nature 387:231–2.
Rieseberg LH, Swensen SM. 1996. Conservation genetics of endangered island plants. In: Avise JC, Hamrick JL (eds). Conservation genetics: case histories from nature. New York NY: Chapman & Hall. p 305–34.
Smith TB, Wayne RK (eds). 1996. Molecular genetic approaches in conservation. New York, NY: Oxford Univ Pr. 483 p.
Taberlet P. 1996. The use of mitochondrial DNA Control region sequencing in conservation genetics. In: Smith TB, Wayne RK (eds.). Molecular genetic approaches in conservation. New York NY: Oxford Univ Pr. p 125–42.
Templeton AR, Shaw K, Routman E, Davis SK. 1990. The genetic consequences of habitat fragmentation. Ann Missouri Botan Garden 77:13–27.
Templeton AR, Georgladis NJ. 1996. A landscape approach in conservation genetics: conserving evolutionary processes in the African Bovidae. In: Avise ]C, Hamrick JL (eds). Conservation genetics: case histories from nature. New York NY: Chapman & Hall p 398–430.
Valladares-Padua C. 1993. The ecology, behavior and conservation of the BLTs (Leontopithecus chrysopygus Mikan, 1823). Unpublished PhD dissertation. Gainesville FL: Univ Florida.
Vane-Wright RI, Smith CR, Kitching IJ. 1994. Systematic assessment of taxic diversity by summation. In; Forey PL, Humphries C], Vane-Wright RI (eds). Systematics and conservation evaluation, systematics association special volume 50. Oxford UK: Clarendon Pr. p 309–26.
Wade MJ, McCauley DE. 1988. Extinction and recolonization: their effects on the genetic differentiation of local populations. Evolution 42:995–1005.
Walker CW. 1997. Patterns of genetic variation in ocelot (Leopardus pardalis) populations for south Texas and northern Mexico. Unpublished PhD Thesis. College Station TX: Texas A&M Univ.
Wang W, Forstner MRJ, Zhang Y, Liu Z, Wei Y, Hu H, Xie Y, Wu D, Melnick DJ. 1997. A phylogeny of Chinese leaf monkeys using mitochondrial ND3–ND4 gene sequences. Intl J Primat 18:305–20.
Wayne RK. 1996. Conservation genetics in the Canidae. In: Avise JC, Hanuick JL (eds). Conservation genetics: case histories from nature. New York NY: Chapman & Hall. p 75–118.
Williams PH, Humphries CJ. 1994. Biodiversity, taxonomic relatedness, and endemism in conservation. In: Forey PL, Humphries CJ, Vane-Wright RI (eds). Systematics and conservation evaluation, systematics association special volume 50. Oxford UK: Clarendon Pr. p 269–87.
Witting L, McCarthy MA, Loeschcke V. 1994. Multi-species risk analysis, species evaluation and biodiversity conservation. In: Loeschcke V, Tomiuk J, Jain SK (eds). Conservation genetics. Basel Germany: Birkhduser Verlag. p 239–49.
Application of Geospatial Information for Identifying Priority Areas for Biodiversity Conservation
Biological diversity, the variety and variability among living organisms and the environment in which they occur, is important to maintain life-sustaining systems of the biosphere, but it is threatened by many human activities. Recently, the United Nations Environment Programme (UNEP) Global Biodiversity Assessment concluded that “the adverse effects of human impacts on biodiversity are increasing dramatically and threatening the very foundation of sustainable development”(UNEP 1995). The total number of species that inhabit the planet is unknown, and most extinctions occur before the species have been named and described. It is estimated that 85–90% of all species can be protected by setting aside areas of high biodiversity before they are further degraded, without the need to inventory species individually. It is generally assumed that most terrestrial species are in the tropics. Realistically, only a relatively small portion of the total tropical land area is likely to be devoted to biodiversity conservation, so it is critical to identify areas rich in species diversity and endemism (the characteristics of species that are native or confined to a particular area) as a first step toward protection of remaining natural habitat before the areas are destroyed.
In the past, protected areas were often set aside without regard to the biodiversity within their boundaries. As a result, many protected areas now have little importance with respect to biodiversity; conversely, many areas of habitat with important biodiversity lack protection. The study discussed here seeks to identify relationships between land cover, human population, and protected areas
through the analysis of comprehensive and consistent spatial datasets at 1-km resolution to answer the following two questions: Are African ecoregions with a high degree of biodiversity adequately protected? Is biodiversity within Africa threatened by human population pressure and land use?
The Study Area
The present study dealt with two areas: the continental area consisted of the African continent, including Madagascar; and the regional area consisted of the African Great Lakes Region, including Burundi, Democratic Republic of Congo, Kenya, Malawi, Mozambique, Rwanda, Tanzania, Uganda, Zambia, and Zimbabwe.
The analysis was carried out with geographic information systems, remote-sensing technologies, and the most comprehensive and consistent 1-km spatial datasets. The land-cover dataset was derived from the International GeosphereBiosphere Program land-cover classification, which was based on the National Oceanic and Atmospheric Administration's 1-km Advanced Very High Resolution Radiometer satellite data spanning a 12-month period (April 1992March 1993). The land-cover characteristic database was produced at the US Geological Survey EROS Data Center. Political boundary data were from the Digital Chart of the World. The protected-areas database from the World Conservation Monitoring Centre and World Resources Africa Data Sampler and the population-density database for Africa from the UNEP Global Resource Information Database were used in the analysis.
Some of the smaller protected areas might not have been accounted for, because of the coarse resolution of analysis. The protected-area database is not current for all countries. The land-cover and population datasets were the best available ones covering all of Africa. Considerable errors are known to exist in the mapped distribution of croplands. The population dataset is generated with a model incorporating many variables, including the location of protected areas, so the areas of intersection between population and protected areas are compromised. However, that does not invalidate conclusions drawn from the analysis regarding the proximity of the protected areas to the areas of high population. None of the datasets has been rigorously validated, so local relationships and distributions should be viewed with caution. Availability of high-quality current data remains a stubborn barrier in such analytical analysis, and this highlights the need to support development and updating of databases.
The Continental Area
Protected areas in Africa occupy slightly over 2 million square kilometers or 7% of the continent's 30 million square kilometers (figure 1). Among various ecoregions, barren and sparsely vegetated lands make up about 9.6 million square
kilometers, whereas biodiversity-rich, tropical evergreen broadleaf forests make up about 3 million square kilometers. Of the barren and sparsely vegetated lands, about 4% of discrete pieces of land are protected, whereas less than 6% of the tropical evergreen broadleaf forests are protected. Closed shrublands, which are estimated to be over 700,000 square kilometers in extent, have the largest proportion of protected area, namely 14%. About 2 million square kilometers, or about 8%, of croplands and a mosaic of croplands mixed with natural vegetation are under protected status.
The Regional Area
The 10-country African Great Lakes Region contains a wide range of habitats, including deserts, savannas, and dry and humid tropical forests. In this region of 6 million square kilometers, 12% of the area is protected. Biodiversity-rich, tropical evergreen broadleaf forests cover about 1.4 million square kilometers of the region, and about 100,000 square kilometers, or slightly less than 7%, is protected, leaving the bulk of the tropical evergreen broadleaf forest unprotected. In contrast, protected areas make up about 9–15% of the areas in the category of woody savannas, savannas, grasslands, croplands, and croplands-natural vegetation mosaic.
Furthermore, the degree to which the forests listed as protected are actually protected varies. In the African Great Lakes Region, for example, about 125,000 square kilometers of croplands and croplands interspersed with natural vegetation mosaic is found in protected areas. This apparent encroachment of agriculture highlights the lack of enforcement of protection of the natural flora and fauna in designated protected areas in the region.
The highest human population densities are found in Rwanda, Burundi, and Uganda around Lake Victoria and in scattered areas in Malawi, Zambia, and Kenya. Areas of low population density coincide with many protected areas, and smaller areas of medium and high population density are found in and adjacent to protected areas.
Summary for Policy-Makers
The geographic analysis of relationships between protected areas, distribution of land-cover types, and population density clearly revealed the following:
• Lack of protection status and effective implementation of protection measures in the designated protected areas seems to pose a serious threat to forest biodiversity in Africa.
• As estimated with a geographic information system, about 7% of the total land area of Africa is protected; this is much higher than the estimate of about 5%, compiled from official statistics, usually cited in international sources. Thus, there are substantial differences between protected-area statistics derived from actual planar area on the ground, as estimated by calculations of a geographic information system, and estimates based on official statistics. The differences, reflecting data of different sources, highlight the need to provide more resources to improve the environmental-information infrastructure in countries so that accurate and up-to-date environmental data can be generated and maintained for planning and policy purposes.
• About 6% of the area covered by biodiversity-rich, tropical evergreen broadleaf forests in Africa is protected. Most of these valuable ecoregions, rich in biodiversity and endemic species, are concentrated in countries like the Democratic Republic of Congo and Madagascar, and seem to lack adequate protection. Practical action programs that include accelerated establishment of a network of protected areas are needed urgently.
• In Africa, drier ecoregions are generally better protected than tropical evergreen broadleaf forests. That is contrary to the widely held belief that moist habitat, such as tropical rain forests, is generally better protected than drier zones, such as dry forests and grasslands.
• The presence of croplands in protected areas indicates that legal designation of areas as protected is not sufficient for the protection of biodiversity in the face of human competition for the same land. Protected status must be accompanied by effective enforcement measures over the long term to ensure protection of biodiversity and endemic and endangered species. Additional resources should be applied to understand socioeconomic factors associated with protection of biodiversity, and local stakeholders should be included by giving them a role and economic incentives to conserve biodiversity.
• In contrast with many other regions, low human population densities in many areas of Africa provide an opportunity to protect such areas for conservation purposes.
• A shift in national and international policy formulation and planning processes based on targeting biodiversity-rich areas is needed to protect biodiversity in Africa more effectively. Geographic targeting and programmatic focus are needed to conserve species ecoregions rich in biodiversity and endemism and to address the socioeconomic causes of encroachment and loss of biodiversity.
The views expressed in this text do not necessarily reflect those of the agencies cooperating in this project. The designations used and material presented above do not imply the expression of any opinion whatsoever on the part of the cooperating agencies concerning the legal status of any country, territory, city, or area or of its authorities or of the delineation of its frontiers or boundaries. The work was jointly funded by the United Nations Environment Programme, the US National Aeronautics and Space Administration, and the US Geological Survey.
My sincere thanks to a number of scientists, including Bhaska Ramachandran, Gene Fosnight, Tom Crawford, Grey Tappan, Brad Reed, Eric Wood, Jim Rowland, Steve Howard of Hughes STX, and Anna Stabrawa of UNEP, who contributed to this work and made valuable suggestions.
UNEP [United Nations Environment Programme]. 1995. Heywood VH (ed.). Global biodiversity assessment. New York NY: Cambridge Univ Pr. 1140 p.
Hawaii Biological Survey:
Museum Resources in Support of Conservation
Hawaiibecause of its geographic isolation, rich volcanic soils, and enormous topographic and climatic diversityhas produced a biota with a very high percentage of endemism among multicellular terrestrial organisms. The native biota includes about 18,000 species (Eldredge and Miller 1998) (table 1). The 8,500 terrestrial and aquatic plants and animals might have evolved from as few as 1,000 original colonists (Gagné 1988; see also Sakai, and others 1995) in the absence of many biotic influences that are present on larger land masses (such as grazing herbivores), and they have proved vulnerable to extreme population reduction and even extinction owing to introduced predators, competitors, and diseases. Although Hawaii accounts for only about 0.2% of the land area of the United States, it has 31% of the nation's endangered species and 42% of its endangered birds. Of the 1,023 species of native flowering plants 73 are down to about 20 or fewer individuals in the wild, and nine are down to one (US Fish and Wildlife Service 1999). Almost 75% of the historically documented extinctions of plants and animals in the United States have occurred in Hawaii.
About 15 years ago, as the dimensions of this extinction crisis were beginning to become clear, a wide array of state, federal, and private organizations, catalyzed by The Nature Conservancy and the Hawaii Audubon Society, redoubled their efforts to develop effective mitigative measures. More recently, a formal consortium of agencies developed the Hawaii Conservation Biology Secretariat, which has raised the profile of these important issues and helped to coordinate responses.
Those efforts have been seriously hampered by lack of fundamental information. The basic taxonomy of many groups has not been fully worked out, and information on the ranges or identities of many species was until recently available only from scattered research publications or museum collections. Although a substantial amount of information has been assembled on endangered plants, vertebrates, and a few invertebrate taxa, successful efforts to manage Hawaiian ecosystems requires information about all species, native and alien. In fact, the greatest threat to Hawaiian organisms and to the integrity of Hawaiian ecosystems is posed by alien species. To address the information need, the Hawaii legislature in 1992 designated the Bishop Museum, which houses the world's largest natural-history collections from Hawaii (nearly 4 million specimens) as the Hawaii Biological Survey (HBS) and charged it with the task of compiling comprehensive information on the entire biota of the state (Allison and others 1995).
The Bishop Museum developed a six-stage process to implement the biological survey. Briefly, this involves, for each major group of organisms,
• developing a computerized database of the literature;
• preparing a species checklist based on the literature, collections, and consultation with experts;
• developing a database of the collections, including coding localities to facilitate geographic information system (GIS) analysis and presentation;
• developing a database of information from other collections or from other organizations that are conducting biological surveys (or establishing computer linkage to such information);
• directing research efforts to high-priority needs; and
• filling gaps in information through additional field surveys.
In practice, many of these are concurrent activities. The literature databases and species checklists developed by HBS scientists and collaborators provide a firm foundation for the computerization of specimen-based data from collections. When specimen data are computerized and incorporated into an environmental information system, one can easily determine the range of a species, document how it has changed, identify broad multispecies patterns of distribution and diversity (ecosystem characteristics), and evaluate how these features are related to various environmental factors (such as climate and soils) and have been or are likely to be affected by resource-management and land-use strategies. It is important to emphasize that specimen collections constitute the most accessible and cost-effective source of data for the development of comprehensive environmental-information systems (Allison 1991; Nielsen and West 1994). Those information systems, involving GIS and other spatial-analysis and database technology, are crucial to the efficient management of Hawaii's fragile ecosystems and are in use by all the state's natural-resource management and land-use agencies.
In its role as HBS, the museum is providing a service to the scientific and local communities as an information clearinghouse. It gathers, processes, synthesizes, and distributes to a variety of partners information related to the biological resources of Hawaii. Information from the collections is crucial to provide authority files, data points for distribution maps, additional ecological information, and a historical perspective on the biota of Hawaii. Inasmuch as completeness is necessary for functionality, HBS also plays a crucial role in centralizing and facilitating distribution of information from partner organizations. The overall strategy is to streamline the process of developing information products while continuing the development of longer-term projects and continuously improving and refining all products.
In this paper, we discuss the overall strategy of HBS and its accomplishments to date. Although our efforts arose out of an urgent need to address critical conservation issues in a relatively small geographic area, we feel that they can serve as an effective model for the role that museums can play in understanding and managing biodiversity. Our overall theme is that museum collections and associated databases are crucial information resources for understanding and managing biological diversity. With more than 400 million specimens in US museums alone, and perhaps 2 billion museum specimens worldwide (Duckworth and others 1993), the implications are enormous.
The information-management strategy developed for HBS is represented schematically in figure 1. Information sources for HBS include those listed on the left
side, including biological surveys, research projects, existing collections, existing literature, and special projects (such as syntheses undertaken with collaborators). In many cases, the flow of information is reciprocal; this is especially true for the collections, where there is constant interaction between scientists producing reports based on the collections, which result in improved quality of identifications and localization. HBS activities are undertaken in collaboration with an array of partner organizations. The collaboration in some cases is formalized at an institutional level, and an informal network of collaboration by scientific staff extends internationally, especially in systematics research. HBS information-based products are used by government, commercial, and private clients for a variety of purposes, including agriculture, conservation, education, fisheries, forestry, health services, land management, quarantine and regulatory services, and other research, as shown on the right side of figure 1.
Some of the primary partners of HBS in recent years have been state and federal natural-resource management agencies (Hawaii Department of Agriculture, Hawaii Department of Land and Natural Resources, US Department of Agriculture, and US Department of the Interior, especially the Fish and Wildlife Service and the former National Biological Survey, now part of the US Geological Survey), conservation organizations (Center for Plant Conservation, Ducks Unlimited, Hawaii Conservation Biology Forum, and The Nature Conservancy), educational organizations (Hawaii Department of Education and University of Hawaii), and other biodiversity research organizations (including Cornell University,
National Tropical Botanic Garden, New York Botanic Garden, and Smithsonian Institution). Many of these organizations maintain specialized databases related to specific applications in conservation or agriculture or to specific taxonomic groups. Rather than duplicate these efforts, we seek to link with them through the development of authority files, data standards, and information models (http://www.bishop.hawaii.org/asc-cnc/).
During the last 5 years, HBS has developed comprehensive bibliographies and species checklists of all major groups of plants and animals and some fungi, protists, and algae in Hawaiiterrestrial, freshwater, and marine. Hawaii is the only state in the United States other than Illinois (Post 1991) and the only large tropical area in the world in which the total number of described species is accurately known (Eldredge and Miller 1995, 1997, 1998; Miller and Eldredge 1996; http://www.bishop.hawaii.org/bishop/HBS/hispp.html).
HBS provides a venue for disseminating work of individual scientists to a variety of users. Most individual researchers do not have at their disposal the contacts, time, or technology needed to deliver their products to all potential users, especially land managers. A researcher might be the world's expert on a particular taxon that occurs in Hawaii but have neither the time nor the means to circulate research results widely within the state. HBS provides an efficient and cost-effective means of disseminating varied research products and extending the useful life of datasets beyond the funding of a particular project or the career of an individual investigator (for example, Helly and others, 1996; US National Committee for CODATA 1995; http://www.sdsc.edu/compeco_workshop/report/helly_publication.html).
The products of HBS take various forms to meet our diverse user community, as shown in figure 2. We see our primary product as information on our World Wide Web (WWW) server. The WWW server makes large amounts of information available worldwide 24 hours a day, and we can update or post information immediately at low cost. Information on the WWW should be our most recent version and should end confusion about versions of information distributed in other media or the use of outdated information that might have been gathered from our collections years ago. The “self-serve” approach also lowers our personnel costs in handling frequently asked questions. Other products beyond the WWW include information services provided directly by staff, enhancements of collections (for example, returning improved identifications of specimens), such technical publications as checklists (Cowie and others 1995; Nishida 1997) and systematic monographs (Gagné 1997), popular publications like our nascent series of user-friendly identification handbooks (Polhemus and Asquith 1996), contributions to formal and informal education, exhibits and internships, and products developed from various partnerships.
One of the products of HBS is the annual publication of a compilation of changes in our understanding of the status and distribution of the Hawaiian biota
titled Records of the Hawaii Biological Survey. Records is published annually in the journal Bishop Museum Occasional Papers. It has been especially effective at providing a publishing vehicle for short papers to document distribution or taxonomic changes that are important in the Hawaii context but might not have an appropriate venue elsewhere in the scientific literature. A number of agencies use the information from HBS to support their own products. One, the Hawaii Ecosystems at Risk project, a consortium led by the US Geological Survey Biological Resources Division, depends on Records as its primary source of documentation of new records of weeds and of taxonomic validation of these records.
We have largely completed the first two of the three levels of databases that provide the foundation for HBS. The first is literature databases. These focus on the taxonomic and distributional literature but include any other publications and reports that come to our attention. The second is taxonomic authority files or species checklists. These databases, compiled largely from the literature with extensive consultation with specialists, provide an index to and synthesis of what has been learned in over 100 years of biological research on Hawaii; without them, much historical information would remain unrecognized or inaccessible. The third is databases of Hawaiian specimens in the Bishop Museum's extensive collections (table 2). Progress in each category of database for each taxon depends on the level of knowledge of the taxon, the expertise available to help, funding priorities, and the curatorial condition of our collections.
Biological surveys are fundamental to the documentation of the plants and animals of the earth (Blackmore and others 1997) and are one of the major reasons for the founding of the world's great natural-history museums (Cotterill 1997; Lane 1996; Raven and others 1993). Early biological surveys were closely associated with exploration of the earth during the last three centuries and had as their purpose documentation of the general biota of scientifically unexplored areas (see, for example, Viola and Margolis 1985). As major biological features of Earth became known, museums' scientific interest shifted more toward detailed taxonomic studies of plant and animal groups. Government agencies were formed to manage natural resources, and they have conducted much of the biological survey work during the last century; for example, in 1939, the Bureau of Biological Survey, an agency in the US Department of Agriculture that developed in close association with the Smithsonian Institution, was, with the Bureau of Fisheries, transferred to the Department of the Interior and later became the Fish and Wildlife Service). With rising human populations and increasing demand for land and natural resources, public and private agencies are now facing tremendous challenges in their efforts to obtain sufficient information to manage and preserve the world's biodiversity.
With the advent of modern database technology, the information in museum collections can be made available for a wide range of uses. This has led to the development of new and strengthened partnerships between museums and resource-management agencies, for example, creation of the National Biological Survey in 1993. These partnerships have focused mostly on the need for detailed information on the distribution of plants and animals to support management
efforts. Museums are the primary repositories of such information. For example, although the systematics of vascular plants of the United States is reasonably well known, precise distributional details on many species are not readily available, and many of the data reside in museum collections; it is therefore urgent to mobilize information from museum collections into databases and to link the databases into information systems.
The major strength of HBS is its comprehensive approach and the fact that its activities are undertaken in close partnership with management agencies. This helps to ensure that HBS products and services meet user needs. In addition, working with partners helps to ensure that collections are built in a purposeful way (see Hawks worth 1991) and have maximal utility. We have emphasized conservation applications in this paper, but biological surveys also have important applications in agriculture, medicine, and recreation (Klassen 1986; Roberts 1992).
The approach of HBS is unique in attempting to provide at least basic information on all organisms while focusing more detailed surveys or products on taxa of concern to specific users. The All Taxa Biodiversity Inventory (ATBI) approach is similar in covering all organisms (Miller 1993; Yoon 1993), but the approaches differ in that HBS synthesizes the literature first and then undertakes surveys to update data and fill gaps, whereas the ATBI emphasizes intensive surveys in smaller areas.
Although many conservation agencies are moving away from efforts to protect individual species and are instead highlighting the need to protect entire ecosystems (Kirlin and others 1994), the classification of ecosystems tends to be rather arbitrary. In a promising alternative approach that has been recently developed (Kiester and others 1996; White and others 1997), species occurrence data (presence or absence) are assembled into map layers and grouped into classes. This method, which can readily use museum-specimen data, involves a high level of objectivity and therefore has many advantages, particularly in public-policy debates, over the use of classed data, such as on vegetation. A particular strength of this approach is that it facilitates analysis and modeling of the risk to biodiversity, including individual species and populations, posed by different landuse strategies.
The scientific importance of museum collections has been well documented (Nudds and Pettitt 1997), but this value is poorly reflected in public policy. Indeed, most museums initially began computerizing their collections to gain internal management efficiency and have been slow to develop scientific products and services outside the traditional research enterprise. The systematics community has also been slow in providing authority files in readily accessible forms, although the recent production of a checklist of almost 100,000 species of North American insects shows what can be done (Poole and Gentili 1997). We agree with Lane (1996) that computerization of collections is central to an expanded role for museums in serving science and society, and nowhere is that more urgent than in the conservation of biodiversity. New organizations throughout the worldsuch as INBio, ERIN, and CONABIO (Anonymous 1994; Gámez 1991; Soberón and others 1996)and long-established organizations, such as the Illinois Natural
History Survey (Anonymous 1996), have proved the importance of museum collections for understanding and managing biodiversity. The recent formation of the US Organization for Biodiversity Information (USOBI) signifies a trend to unite individual institutional efforts into a federation to achieve economies of scale and develop standards and common gateways to highly dispersed data (NRC 1993:94–5).
We thank our many collaborating individuals and institutions, especially the John D. and Catherine T. MacArthur Foundation and the National Science Foundation for major funding. Gordon Nishida prepared the figures.
Allison A. 1991. The role of museums and zoos in conserving biological diversity in Papua New Guinea. In: Pearl M, Beehler BM, Allison A, and Taylor M (eds), Conservation and environment in Papua New Guinea: establishing research priorities. Washington DC: The Government of Papua New Guinea and Wildlife Conservation International. p 59–63.
Allison A, Miller SE, and Nishida GM. 1995. Hawaii Biological Surveya model for the Pacific Region. In: Maragos JE, Peterson MNA, Eldredge LG, Bardach JE, and Takeuchi HF (eds). Marine and coastal biodiversity in the tropical island Pacific region. Vol 1, p 349–55. Honolulu HI: East-West Center.
Anonymous. 1994. Collaboration with biodiversity agencies. Erinyes 20:1–8.
Anonymous. 1996. Illinois Natural History Survey Annual Report 1995–1996. Champaign IL: Illinois Natural History Survey, 55 p.
Blackmore S, Donlon N, and Watson E. 1997. Calculating the financial value of systematic biology collections. In: Nudds JR, and Pettitt CW (eds). The value and valuation of natural science collections. Proceedings of the International Conference, 1995, Manchester UK. London UK: Geological Society. p 17–21.
Cotterill FPD. 1997. The second Alexandrian tragedy and the fundamental relationship between biological collections and scientific knowledge. In Nudds, JR & Pettit, CW (eds), Proceedings of the International Conference on the Values and Valuation of Natural Science Collections. p. 227–241. Manchester: Manchester Museum.
Cowie RH, Evenhuis NL, Christensen CC. 1995. Catalog of the native land and freshwater molluscs of Hawaii. Leiden Netherlands: Backhuys Publ. 248 p.
Duckworth WD, Genoways HH, Rose CL. 1993. Preserving natural science collections: chronicle of our environmental heritage. Washington DC: National Institute for the Preservation of Cultural Property Inc. 140 p.
Eldredge LG, Miller SE. 1995. Records of the Hawaii Biological Survey for 1994. How many species are there in Hawaii? Bishop Mus Occas Pap 41:1–18.
Eldredge LG, Miller SE. 1997. Numbers of Hawaiian species: supplement 2, including a review of freshwater invertebrates. Bishop Mus Occas Pap 48:3–22.
Eldredge LG, Miller SE. 1998. Numbers of Hawaiian species: supplement 3, with notes on fossil species. Bishop Mus Occas Pap 55:3–15.
Gagné WC. 1988. Conservation priorities in Hawaiian natural systems. BioScience 38:264–71.
Gagné WC. 1997. Insular evolution, speciation, and revision of the Hawaiian genus Nesiomiris (Hemiptera: Miridae). Bishop Mus Bull Entomol 7: i–x, p 1–226.
Gámez R. 1991. Biodiversity conservation through facilitation of its sustainable use: Costa Rica's National Biodiversity Institute. Trends Ecol Evol 6(12):377–8.
Hawksworth DL (ed). 1991. Improving the stability of names: needs and options. Regnum Vegetabile No. 123. Königstein : Koeltz Scientific Bk.
Helly JT, Case T, Davis F, Levin S, Michener W(eds.). 1996. The state of computational ecology. San Diego CA: San Diego Supercomputer Center. 20pp. [also at http://www.sdsc.edu/compeco_workshop/report/helly_publication.html.
Kiester AR, Scott JM, Csuti B, Noss RF, Butterfield B, Sahr K, and White D. 1996. Conservation priorization using GAP data. Cons Biol 10(5):1332–42.
Kirlin JJ, Asmus P, Thompson R. 1994. Species conservation through ecosystem management. California Policy Choices 9:143–171.
Klassen W. 1986. Agricultural research: the importance of a national biological survey to food production. In: Kim KC Knutson L (eds). Foundations for a national biological survey. Lawrence KS: Assoc of Systematics Collections. p 65–76.
Lane MA. 1996. Roles of natural history collections. Ann Missouri Botan Gard 83:536–45.
Miller SE. 1993. All Taxa Biological Inventory workshop. Assoc Syst Coll News 21(4): 41, 46–7.
Miller SE, Eldredge LG. 1996. Number of Hawaiian species: supplement 1. Bishop Mus Occais Pap 45:8–17.
Mlot C. 1995. In Hawaii, taking inventory of a biological hot spot. Science 269:322–3.
NRC [National Research Council]. 1993. A biological survey for the nation. Washington DC: NatlAcad Pr. 205 p.
Nielsen ES, West JG. 1994. Biodiversity research and biological collections: transfer of information. In: Forey PL, Humphries CJ, Vane-Wright RI (eds). Systematics and conservation evaluation. Oxford UK: Clarendon Pr. p 101–21.
Nishida GM. 1997. Hawaiian terrestrial arthropod checklist. Third edition. Bishop Mus Tech Rep 12.
Nudds JR, Pettitt CW (eds). 1997. The value and valuation of natural science collections. Proceedings of the International Conference, Manchester, 1995. Manchester UK: Manchester Museum. p xii + 276.
Polhemus D, Asquith A. 1996. Hawaiian damselflies: a field identification guide. Hawaii Biol Surv Handbook. Honolulu HI: Bishop Museum Pr. 122 p.
Poole RW, Gentili P (eds). 1997. Nomina Insecta Nearctica: a check list of the insects of North America. Rockville MD: Entomological Information Services. CD-ROM. Also published as check list in four paper volumes. 1996–1997.
Post SL. 1991. Native Illinois species and related bibliography. Illinois Nat Hist Surv Bull 34:463–75.
Roberts L. 1992. Chemical prospecting: hope for vanishing ecosystems. Science 256:1142–3.
Sakai AK, Wagner WL, Ferguson DM, Herbst DR. 1995. Origins of dioecy in the Hawaiian flora. Ecology 76:2517–29.
Soberón J, Llorente J, Benítez H. 1996. An international view of national biological surveys. Ann Missouri Botan Gard 83:562–73.
US Fish and Wildlife Service. 1999. US Fish and Wildlife Service Species List, March 23, 1999. Honolulu: unpubl.
US National Committee for CODATA, Committee for Pilot Study on Database Interfaces. 1995. Finding the forest in the trees: the challenge of combining diverse environmental data: selected case studies. Washington DC: Natl Acad Pr. 129 p.
Viola HJ, Margolis C. 1985. Magnificent voyagers: the US Exploring Expedition, 1838–1842. Washington DC: Smithsonian Inst Pr. 303p.
White D, Minotti PG, Barczak MJ, Sifneos JC, Freemark KE, Santelmann MV, Steinitz CF, Kiester AR, Preston EM. 1997. Assessing risks to biodiversity from future landscape change. Cons Biol 11(2):249–360.
Yoon CK. 1993. Counting creatures great and small. Science 260:620–2.
Building the Next-Generation Biological-Information Infrastructure.
A grand challenge for the 21st century is to harness the accumulating knowledge of Earth's biodiversity and the ecosystems that support it. To accomplish that, we must mobilize biological informationassemble it, organize it, and deliver it with dramatically increased capacity. We must elevate the global biological-information infrastructure to a new level of capabilitya “next generation”that will allow people to share on a worldwide basis the knowledge created by biodiversity and ecosystems research.
Recognizing the urgency of the task, the President's Committee of Advisors on Science and Technology, through its Panel on Biodiversity and Ecosystems, recently coordinated a review of the US National Biological Information Infrastructure (NBII) (PCAST 1998). Over a 6-month period in 1997, people from a broad cross section of the public and private sectors contributed their insights, experiences, concerns, and hopes. What emerged was a renewed understanding of the importance of biological information to all aspects of human society. It became clear that much remains to be done to ensure that this information is complete and usable. Although the purpose of the review was to develop recommendations to build capacity in the United States, many of the panel's findings address global concerns of relevance to biodiversity research wherever it occurs. In this paper, we provide a summary of the panel's report, a view of what a next-generation
biological-information infrastructure might encompass, and suggestions about how it might be achieved.
In the United States, NBII is the primary mechanism whereby biodiversity and ecosystem information is made available to all sectors of society. It is the biological component of the National Information Infrastructure and is the framework that connects US activities to the global biodiversity and ecosystem research enterprise. Its meaning is expansive and intended to convey the idea that an information infrastructure comprises not only computers, networks, and the like, but also the information, policies, standards, and people who use it. Initiation of the NBII was one of the primary recommendations made by the 1993 National Research Council report A Biological Survey for the Nation (NRC 1993).
Because our fate and economic prosperity are so completely linked to the natural world, information about biodiversity and ecosystemsas well as the infrastructure that supports itis vital to a wide range of scientific, educational, commercial, and government uses. Most of this information now exists in forms that are not easily accessed or used. From traditional paper-based libraries to scattered databases and physical specimens preserved in natural-history collections throughout the world, our record of biodiversity and ecosystem resources is uncoordinated, and large parts of it are isolated from general use. It is not being used effectively by scientists, resource mangers, policy-makers, or other potential client communities (National Performance Review 1997; NRC 1997).
Research activities are being conducted around the world that could improve our ability to manage biological information. In the United States, the Human Genome Project is producing new medical therapies and developments in computer and information science. Geographic information systems (GISs) are expanding the ability of federal agencies to conduct data-gathering and data-synthesis activities more responsibly and creating opportunities for commercial partnerships that can lead to new software tools. The National Spatial Data Infrastructure (http://nsdi.usgs.gov) is improving the management of geographic, geological, and satellite datasets; the Digital Libraries (http://www.cise.nsf.gov/iis/dli/home.html) projects are beginning to produce useful results for some information domains; and the High-Performance Computing and Communications Initiative (http://www.hpcc.gov) has enhanced some computation-intensive engineering and science fields.
But little attention has been paid to computer and information science and technology research in the biodiversity and ecosystem domain. We must produce mechanisms that can efficiently search through terabytes of Mission to Planet Earth satellite data and other biodiversity and ecosystem datasets, make correlations among data from disparate sources, compile those data in new ways, analyze and synthesize them, and present the results in an understandable and usable manner. Despite encouraging advances in computation and communication performance in recent years, we are able to perform these activities on only a very small scale. We can, however, make rapid progress if the computer and
information science and technology research community becomes focused on the needs of the biodiversity and ecosystem research community (Robbins 1996).
Knowledge about biodiversity and ecosystems is vast and complex. The complexity arises from two sources. The first is the underlying biological complexity of the organisms themselves. There are millions of species, each of which is highly variable across individual organisms, populations, and time. Species have complex chemistries, physiologies, developmental cycles, and behaviors resulting from more than 3 billion years of evolution. There are hundreds, if not thousands, of ecosystems, each comprising complex interaction among large numbers of species and between those species and multiple abiotic factors.
The second source of complexity in biodiversity and ecosystem information is sociologically generated. The sociological complexity includes problems of communication and coordinationamong agencies, among divergent interests, and among groups of people from different regions and different backgrounds (academe, industry, and government) and with different views and requirements. The kinds of data that humans have collected about organisms and their relationships vary in precision, in accuracy, and in numerous other ways. Biodiversity data types include text and numerical measurements, images, sound, and video. The range of other databases with which biodiversity datasets must interact is also broad, including geographic, meteorological, geological, chemical, and physical databases. The mechanisms used to collect and store biological data are almost as varied as the natural world that they document. In addition, biological data can be politically and commercially sensitive and can entail conflicts of interest. Users' skill levels are highly variable, and training in this field is not well developed.
Because of those complexities, humans still play a crucial role in the processing of biological data. Biological information is not as amenable to automatic correlation, analysis, synthesis, and presentation as many other types of information, such as that in radioastronomy, where there is more coherent global organization and the problems being studied are often conducive to automatic analysis. In biodiversity research, people act as sophisticated filters and query processorslocating resources on the Internet, downloading datasets, reformatting and organizing data for input to analysis tools, then reformatting again to visualize results. This process of creating higher-order understanding from dispersed datasets is a fundamental intellectual process, but it breaks down quickly as the volume and dimensionality of the data increase. Who could be expected to “understand” millions of cases, each having hundreds of attributes? Yet problems on this scale are common in biodiversity and ecosystem research (Schnase and others 1997).
For a biological-information infrastructure to be effective, it must provide the means to manage complexity. It must allow scientists to extract new knowledge from the aggregate mass of information generated by the data-gathering and synthesis activities of other scientists. It must use the power of computers to facilitate the queries, correlations, and processing that are impossible for humans to
perform alone. And it must deliver this functionality within a physically and intellectually accessible framework. That means developing ways of delivering information to a wide array of users with differing skills, ages, and investment in the material.
We are only beginning to develop a vocabulary to describe these large-scale, synthetic, information-processing activities. Some sociologists use the term distributed cognitive system to emphasize the role of humans in a synergistic information-processing network (Hutchins 1995). Data mining is often used by the data base community. Whatever the name, the activities form only a part of a process of knowledge discovery that includes the large-scale, interactive storage of information (known by the unintentionally uninspiring term data warehousing); the cataloging, cleaning, preprocessing, transformation, verification, and reduction of data; and the generation and use of models, evaluation and interpretation, personal communication, the evolution of sophisticated user interfaces, and finally consolidation and use of the newly extracted knowledge. Those processes will become increasingly important if we are to use what we know and expand our knowledge in useful directions.
At present, the NBII provides little support for these activities. At best, it can be used to access information in databases held by federal agencies and other institutions around the country. Once the information is accessed, however, the task of organizing, integrating, and interpreting it remains, for the most part, a laborious, manual process. The development of computational tools for the biodiversity and ecosystem enterprise lags behind other sciences. Important classes of information are missing (information on fewer than 1% of the specimens in our natural-history collections has been entered in databases!), and databases are uneven in the types of information that they hold. It is difficult for individual scientists to publish their data electronically in useful ways. Standards for information exchange have not been widely adopted. We have no mechanism for archiving data over generations of use and generations of technologies. And the power of communication networks to build communities remains largely untapped. In summary, the NBII is neither a system nor an infrastructure: it is a cumbersome and brittle patchwork that presents as many obstacles to scientific work as it does opportunities. It clearly is time to transform it into a coherent and empowering capability.
The Next GenerationNBII-2
We envision a “next generation” National Biological Information Infrastructure, NBII-2, that would address many of the concerns described above. The overarching goal of NBII-2 would be to become a fully accessible, distributed, interactive digital library. NBII-2 would provide an organizing framework from which scientists could extract useful informationnew knowledgefrom the aggregate mass of information generated by various data-gathering activities. That would be accomplished by using the power of computers and communication networks to augment the processing activities that now require a human mind. It would make analysis and synthesis of vast amounts of data from multiple datasets easier
and more accessible to a variety of users. It would also serve management and policy decision-making, education, recreation, and industry by presenting data to each user in a manner tailored to that user's needs and skill level.
We envision NBII-2 as a distributed facility that would be considerably different from a “data center,” considerably more functional than a traditional library, considerably more encompassing than a typical research institute. Unlike a data center, NBII-2 would have the objective of automatic discovery, indexing, and linking of datasets rather than collection of all datasets on a given topic into one facility. Following the best practice of traditional libraries, this special library would update the form of storage and upgrade information content as technologies evolve. Unlike a typical research institute, it would provide services to research going on elsewhere, and its own staff would conduct biodiversity and ecosystem research and research in biological informatics. The facility would offer “library” storage and access to diverse constituencies.
The core of NBII-2 would be a “research library system” that would comprise at least five regional nodes sited at appropriate institutions (national laboratories, universities, museums, and so on) and connected to each other and to the nearest telecommunication providers by the highest-bandwidth network available. In addition, NBII-2 would seamlessly integrate all computerslaptops, workstations, fileservers, and supercomputerscapable of storing and serving biodiversity and ecosystem data via the Internet. The providers of information would have complete control over their own data but have the opportunity to benefit from (and the right to refuse) the data-indexing, cleansing, and long-term storage services of the system as a whole.
NBII-2 would be
• the framework to support knowledge discovery for the nation's biodiversity and ecosystem enterprise and to involve many client and potential-client groups;
• a common focus for independent research efforts and a global context for sharing information among those efforts;
• an accrete-only, no-delete facility from which all information would be available on line24 hours a day, 7 days a weekin a variety of formats;
• a facility that would serve the needs of (and eventually be supported by partnership with) government, the private sector, education, and individuals;
• an organized framework for collaboration among federal, regional, state, and local organizations in the public and private sectors that would provide improved programmatic efficiencies and economies of scale through better coordination of efforts;
• a commodity-based infrastructure that uses readily available, off-the-shelf hardware and software and the products of digital-library research wherever possible;
• an electronic facility where scientists and others could “publish” biodiversity and ecosystem information for cataloging, automatic indexing, access, analysis, and dissemination;
• a place where intensive work on how people use large information systems would be conducted, including studies of human-computer interaction, the
sociology of scientific practice, computer-supported cooperative work, and user-interface design;
• a place for developing the organizational and educational infrastructure that will support sharing, use, and coordination of massive datasets;
• a facility that would provide content storage resources, registration of datasets, and “curation” of datasets (including migration, cleansing, and indexing);
• an applied biodiversity and ecosystem informatics research facility that would develop new technologies and offer training in informatics; and
• a facility that would provide high-end computation and communication to researchers and institutions throughout the country.
The facility would not be a purely technical and technological construct but, would also encompass sociological, legal, and economic issues within its research purview. These would include intellectual-property rights management, public access to the scholarly record, and the characteristics of evolving systems in the networked information environment. The human dimensions of the interaction with computers, networks, and information will be particularly important subjects of research as systems are designed for the greatest flexibility and usefulness to people.
The research nodes of NBII-2 must address many needs, including
• new statistical pattern-recognition and modeling techniques that can work with high-dimensional, large-volume data;
• workable data-cleaning methods that automatically correct input and other types of errors in databases;
• strategies for sampling and selecting data;
• algorithms for classification, clustering, dependency analysis, and change and deviation detection that scale to large databases;
• visualization techniques that scale to large and multiple databases;
• metadata encoding routines that will make data mining meaningful when multiple distributed sources are searched;
• methods for improving connectivity of databases, integrating data-mining tools, and developing better synthetic technologies;
• methods for improving large-scale project coordination and scientific collaborations;
• continuing, formative evaluation, detailed user studies, and quick feedback between domain experts, users, developers, and researchers;
• methods for facilitating data entry and the digitization of large amounts of irregularly structured information; and
• ways of engaging society in the pursuit of global information-sharing.
None of those problems is peculiar to biodiversity research. However, there is an urgent need to address them in the biodiversity domain because research has demonstrated that there can be no domain-independent solutions. We cannot “borrow” discoveries wholesale from other disciplines; we must work through these problems ourselves (Star and Ruhleder 1996). To comprehend and use our
biodiversity and ecosystem resources, we must learn how to exploit massive datasets, learn how to store and access them for analytical purposes, and develop methods to cope with growth and change in data. NBII-2 as envisioned here can be the enabling framework that unlocks the knowledge and economic power lying dormant in the masses of biodiversity and ecosystem data that we have on hand now and will accumulate in the future.
The total volume of biodiversity and ecosystem information is almost impossible to measure. We do know that whatever the total, only a fraction has been captured in digital form. Our natural-history museums, for example, contain at least 750 million specimens, the vast majority of which have not been recorded in databases. The same holds for the published record, where most biodiversity and ecosystem information still resides in paper-based journals, books, field notes, and the like. Clearly, one of the most important infrastructure issues is to move the biodiversity and ecosystem enterprise into a digital worldto create the content for the NBII-2 digital libraryby digitizing the existing corpus of scholarly work on a large scale.
The NBII-2 digital library will place challenging demands on network hardware services and on software services related to authentication, integrity, and security. Needed are both a fuller implementation of current technologies, such as digital signatures and a public-key infrastructure for managing cryptographic key distribution, and consideration of tools and services in a broader context related to library use. For example, the library system might have to identify whether a user is a member of an organization that has some set of access rights to an information resource. As a national and international enterprise that serves a large range of users, the library must be designed to detect and adapt to various degrees of accessibility of resources connected to the Internet.
A fully digital, interactive library system, such as NBII-2, will require substantial computational resources, although little is known now about the precise scope of the necessary resources. In many aspects that are critical to digital libraries, such as knowledge representation and resource description or summarization and navigation, even the basic algorithms and approaches are not yet well defined, so it is difficult to project computational requirements. Many information-retrieval techniques are intensive in their computational and input-output demands as they evaluate, structure, and compare large databases in a distributed environment. Distributed-database searching, resource discovery, automatic classification and summarization, visualization, and presentation are also computationally intensive activities that are likely to be common in the NBII-2 digital library.
Finally, NBII-2 will require enormous storage capacity. Even though the library system we are proposing would not set out to accrue datasets to become the repository of all biodiversity datamany other federal agencies have their own storage facilities, and various data-providers will want to retain control over their own datalarge amounts of storage on disk, tape, optical media, and other future storage forms will still be required. As research is conducted to produce new ways to
manipulate large datasets, these will have to be sought out, copied from their original sources, and stored for use in the research. And in serving its long-term curation function, NBII-2 will accumulate substantial amounts of data for which it will be responsible, including redundant datasets that will have to be maintained in case of loss.
New approaches to managing information must be developed in the context of NBII-2. Massive datasets can lead to the collapse of traditional approaches in database management, statistics, pattern recognition, personal-information management, and visualization. For example, a statistical-analysis package assumes that all the data to be analyzed can be loaded into memory and then manipulated. What happens when the dataset does not fit into main memory? What happens if the database is on a remote server and will never permit a naive scan of the data? What happens if queries for stratified samples cannot be accepted, because data fields in the database being accessed are not indexed and the appropriate data therefore cannot be located? What if the database is structured with only sparse relations among tables or if the dataset can be accessed only through a hierarchical set of fields?
Furthermore, challenges often are not restricted to issues of scalability of storage or access. For example, what if a user of a large data repository does not know how to specify the desired query? It is not clear that a structured query language (SQL) statementor even a programcan be written to retrieve the information needed to answer a query like, “Show me the list of gene sequences for which voucher specimens exist in natural-history collections and for which we also know the physiology and ecological associates of those species.” Many of the interesting questions that users of biodiversity and ecosystem information would like to ask are of this type: they are “fuzzy,” the data needed to answer them must come from multiple sources that will be inherently different in structure and conceptually incompatible, and the answers might be approximate.
Major advances are needed in methods for knowledge representation and interchange, database management and federation, navigation, modeling, and data-driven simulation; in approaches to describing large, complex networked information resources; and in techniques to support networked information discovery and retrieval in extremely large-scale distributed systems. In addition to near-term operational solutions, new approaches are needed to longer-term issues, such as the preservation of digital information across generations of storage, processing, and representation technology. Traditional information-science skills, such as thesaurus construction and indexing, must be elaborated on and scaled to accommodate large information sources. We need to preserve and support the knowledge of library-science and information-science researchers and help to scale up the skills of knowledge organization and information retrieval.
Also much needed are software applications that provide more-natural interfaces between humans and databases than are now available. For example, a valuable data-cleansing activity might be to “show the data related to all specimens
in our natural-history collections whose likelihood of being mislabeled exceeds 0.75.” Assuming that some cases in the database can be identified as “labeled correctly” and others “known to be mislabeled,” a training sample for a data-mining algorithm could be constructed. The algorithm would build a predictive model and retrieve records matching that model rather than a structured query that a person might write. This is an example of a much needed and much more natural interface between humans and databases than is currently available. In this case, it eliminates the requirement that the user adapt to the machine's needs rather than the other way around. We must refine and augment the interactions between people and machines, expand the role of agentry in information systems, and discover more-powerful and more-natural ways of navigating the scientific record.
In return, research in computer and information science and technology in the biodiversity and ecosystem domain is likely to yield discoveries of value to other fields (Spasser 1998). Nowhere do we find the problems of heterogeneous database federation more challenging than in the life sciences. A fully implemented digital library for biology would include everything from ideas to physical objects and enormous amounts of information in every medium type imaginable. Research on global climate change, habitat destruction, and the discovery of species is among the most distributed of our scientific activities and creates extraordinary opportunities to learn about computer-mediated project coordination and communication. At almost every turn, scale, complexity, and urgency conspire to create a particularly wicked set of problems. Working on these problems will undoubtedly advance our understanding and use of information technologies, perhaps more than in any other circumstance.
We have laid out the case for building a fully digital, interactive, research-library system for biodiversity and ecosystem information and the basic requirements of and goals for the library and its research and service. But how much will it cost, and how long will it take to build?
We estimate that each of the regional nodes that will form the core of NBII-2 will require an annual operating budget of at least $8 millionprobably more. Minimally, supporting five such nodes would require at least $40 million per year, an amount that is a small fraction of the funds spent nationwide each year to collect data (conservatively estimated at $500 million for federal government projects alone). As with the Internet itself, the federal government should provide the “jump start” for this new infrastructure by investing heavily in its formative stages. Part of the investment should be devoted to developing incentives for the participation of private-sector partners. Gradually, support and operation of the infrastructure should be shared by nongovernment participants, as has happened with the Internet.
The planning and request-for-proposals process should be conducted within a year. Merit review and selection of sites should be complete within the following six months. The staffing of the sites and initial coordination of research and
outreach activities should take no more than a year after initial funding is provided. The “lifetime” of each facility should not be guaranteed for more than 5 years, but the system must be considered a long-term activity so that data access is guaranteed in perpetuity. Evaluation of the sites and of the system should be regular and rigorous, although the milestones whereby success can be measured will be the incremental improvements in ease of use of the system by students, policy-makers, scientists, and others. In addition, an increasing number of public-private partnerships that fund the research and other operations will indicate the usefulness of accessible, integrated information to commercial and government interests.
In the 21st century, work will depend increasingly on rapid, coordinated access to shared information. Through the shared digital library of NBII-2, scientists and policy-makers will be able to collaborate with colleagues who are geographically and temporally distant. They will use the library to catalog and organize information, perform analyses, test hypotheses, make decisions, and discover new ideas. Educators will use its systems to read, write, teach, and learn. In traditional fashion, intellectual work will be shared with others through the medium of the librarybut these contributions and interactions will be elements of a global and universally accessible library that can be used by many different people and many different communities. By increasing the effectiveness of information, NBII-2 is likely to lead to scientific discoveries, advance existing fields of study, promote disciplinary fusions, and enable new research traditions. And most important, it could help us to protect and manage our natural capital so as to provide a stable and prosperous future.
Hutchins E. 1995. Cognition in the wild. Cambridge MA: MIT. 381 p.
National Performance Review. 1997. Access America: reengineering through information technology. Report of the National Performance Review and the Government Information Technology Services Board. Washington DC: GPO. 97 p.
NRC [National Research Council]. 1993. A biological survey for the nation. Washington DC: National Acad Pr. 205 p.
NRC [National Research Council]. 1997. Bits of power: issues in global access to scientific data. Washington DC: National Acad Pr. 235 p.
PCAST [President's Committee of Advisors on Science and Technology]. 1998. Teaming with life: investing in science to understand and use America's living capital. Report to the President of the United States from the PCAST Panel on Biodiversity and Ecosystems. Washington DC: GPO.
Robbins RJ. 1996. Bioinformatics: essential infrastructure for global biology. J Comp Biol 3(4):465–78.
Schnase JL, Kama DL, Tomlinson KL, Sánchez JA, Cunnius EL, Morin NR. 1997. The flora of North America digital library: a case study in biodiversity database publishing. J Network Comp Applica 20:87–103.
Spasser MA. 1998. Articulating collaborative activity: design-in-use of collaborative publishing services in the Flora of North America Project. Proceedings of ISCRAT '98 (Århus, Denmark, June 1998).
Star SL, Ruhleder K. 1996. Steps toward an ecology of infrastructure: design and access for large information spaces. Info Syst Res 7(1):111–34.