Systematics, biological knowledge and environmental ... - Springer Link

Biodiversity and Conservation 4, 183-205 (1995)

Systematics, biological knowledge and environmental conservation F.P.D. C O T T E R I L L Biodiversity Foundation for Africa* Secretariat: PO Box FM730, Famona, Bulawayo, Zimbabwe and Curator of Mammalogy, Natural History Museum of Zimbabwe, PO Box 240, Bulawayo, Zimbabwe

Received 8 June 1994; revised and accepted 12 October 1994

Systematics and taxonomy are essential: they respectively elucidate life's history, and organize and verify biological knowledge. This knowledge is built of interrelated concepts which are ultimately accounted for by biological specimens. Such knowledge is essential to decide how much and what biodiversity survives human onslaughts. The preservation of specimens in natural history collections is the essential part of the process which builds and maintains biological knowledge. These collections and the human expertise essential to interpret specimens are the taxonomic resources which maintain accurate and verifiable concepts of biological entities. Present and future knowledge of the complexities and diversity of the biosphere depends on the integrity of taxonomic resources, yet widespread ignorance and disregard for their fundamental value has created a global crisis. Preservation of specimens in natural history collections is chronically neglected and support to study and manage collections is very insufficient. The knowledge held by experienced taxonomists is not being passed on to younger recruits. Neglect of collections has destroyed countless specimens and threatens millions more. These threats to taxonomic resources not only impinge on systematics but all biology: this tragedy jeopardizes the integrity of biological knowledge. The consequences for environmental conservation and therefore humanity are also of dire severity and the biodiversity crisis adds unprecedented weight to the barely recognized crisis in taxonomy and systematics. Keywords: biological collections; conservation; biodiversity information; phylogenetic systematics;

taxonomy.

Introduction This paper examines the paradoxical plight and importance of systematics and taxonomy in detail and endorses the need for an objective knowledge of biodiversity to support environmental conservation. The interface between the latter and systematics and taxonomy has to do with the ability of these disciplines to improve and organize our knowledge of biological complexity. I argue that the state of taxonomy and systematics directly impinges on the integrity of biological knowledge and the values we place on biodiversity. Biodiversity describes the variety of all life forms and life processes which form the complex ecosystems of the biosphere (Noss, 1990). Because systematics improves and organizes our knowledge of this complexity, the science is vital to the study of biodiversity (Eldredge, 1992). Systematics discovers biological history: understanding the origins and *Where correspondence should be addressed. 0960-3115 © 1995 Chapman & Hall

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relationships among extinct and extant life forms (O'Hara, 1988) and taxonomy is the science of naming and classifying these organisms (de Queiroz and Gauthier, 1994). Taxonomy and systematics are frequently undervalued by society, which is strange because both disciplines are indispensable to any scientific endeavour aimed at elucidating the history of life on earth, and the interrelationships among its components (Gould, 1989). Natural history collections and the human skills to manage and identify the specimens they contain are the key resources of systematics. They are, respectively, the sources and the means to interpret biodiversity. These constitute the taxonomic resources, which underpin scientific efforts to elucidate biological complexity, understand evolutionary history and support environmental conservation. It is paradoxical that their fundamental value is largely unappreciated and poorly supported. Both the scope of existing collections and available taxonomic expertise are insufficient to complete an exploration of the biosphere in reasonable time (Lane, 1992; Raven and Wilson, 1992; May, 1993). The inadequacy of people and resources to carry out the inventory and interpretation of organisms required to counter the biodiversity crisis was originally addressed by Wilson (1985), one of the first scientists to express alarm over the plight of taxonomy (Wilson, 1971) was labelled 'Biological Diversity Crisis II' (Boom, 1991). Worldwide, support for systematics and taxonomic resources is decreasing, despite the increased commitments required (Krebs, 1992; Linnaean, 1992; de la Penha, 1993; Idema, 1993). Although concern over the state of systematics is apparent (Crowe et al., 1989; Harvey, 1991; Krebs, 1992; Lane, 1992; Linnaean, 1992; Alberch, 1993; Parnell, 1993; Rose et al., 1993), the majority of environmentalists and biologists are either uninformed or unconcerned about 'Biodiversity Crisis If'. We cannot disregard two cardinal premises in biology that are critical to sustain and communicate reliable knowledge: biological entities (principally organisms) must be accurately identified and classified in relation to each other. Adherence to these principles not only maintains accurate knowledge, but ensures its universal communication. The sanctioning of these principles will singularly impinge on the future of taxonomic resources and thus the integrity of biological knowledge. A consensus is needed, to acknowledge and respond to a state of affairs where: (1) Taxonomic resources are under burgeoning threats. It is imperative we recognize that losses of specimens destroys the quintessential foundations supporting biological knowledge, and that losses of taxonomic expertise removes the means to decipher and communicate critical information. (2) The scientific processes, whereby correct information on the biosphere is obtained and biological knowledge is accounted for, must become widely understood and supported. It is therefore crucial to inform society that biological collections are sources of scientific information of the natural world, and illuminate how systematics interprets the origins, identities and relationships among organisms. (3) Taxonomic resources, and especially biological collections, are of international worth. Those accountable for biological collections have a concomitant responsibility to maintain them for present and future generations. The vital roles of taxonomic resources in managing complexities of interrelated facts and maintaining the veracities of biological knowledge must be actively marketed to society, scientists and decision makers. The importance of biodiversity can no longer be ignored (Wilson, 1992; May, 1993).

Systematics, biological knowledge and conservation

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Understanding the natural world and halting the biodiversity crisis ultimately depends on a vibrant state of systematics. Sustainable and successful conservation of the environment is ultimately reliant on an efficient infrastructure of taxonomic resources to supply and manage a scientific knowledge of biodiversity (Janzen, 1993a, b). Unfortunately, even the elementary information on which to base objective decisions on environmental issues is invariably not available (Pressey et al., 1993) and such acute deficiencies in our knowledge are especially true of tropical biodiversity (McNeely et al., 1990; Andr6 et al., 1992; Wilson, 1992). Over the past decade, the realization that we need a detailed scientific knowledge of the natural world to halt the environmental crisis (Hardin, 1985), has attracted much attention to systematics (Soul6, 1990; Eldredge, 1992; Linnaean, 1992; Alberch, 1993). We need to be more specific about the details of the scientific knowledge we require of biodiversity. Inadequate support for systematics Most taxonomists currently operate in exile, only working on collections in their spare time '... in large part because they cannot get funding to do taxonomy and build on that base' (Janzen, 1993a). This inability to secure support epitomizes the image problem of systematics (McAlpine, 1986; Crowe et al., 1989) and illustrates society's naivety of the complexities of the natural world, or how science can interpret the apparent obscurities of biodiversity. It appears that many funding agencies, though claiming to support biodiversity conservation, routinely reject proposals with a taxonomic content. This apparently stems from an ignorance of what biodiversity really is and how it is studied. This can no longer be excused, as specific guidelines unequivocally identify and underwrite the need to support systematics in order to conserve biological diversity (NSB, 1989; NSF, 1992). In spite of these guidelines, the articulation of the value of systematics within the context of the biodiversity crisis (Eldredge, 1992; Krebs, 1992; Wilson, 1992), and the considerable attention attracted to these issues since the Earth Summit (Strong, 1993; di Castri, 1993), effective support for systematics remains very inadequate. The performance of the Global Environment Facility (GEF) - the funding mechanism delegated at the Earth Summit to support biodiversity conservation - has been an embarrassing waste of resources, as judged by its mismanagement to date, lack of clear objectives and insufficient funding of deserving projects (Mittermeier and Bowles, 1993). The GEF would greatly benefit biodiversity conservation if it supported the fragmented infrastructure of threatened resources (collections and associated human skills) which represents taxonomy. Mass extinction of taxonomic resources Loss of hurnan expertise As individual systematists retire or die in reaching the end of a lifetime's research, effective transfer of their expertise to new recruits is the exception (Marshall, 1993; Parnell, 1993). Hundreds of lifetimes of experience (typified by the expertise to identify particular groups of organisms efficiently and accurately) is dying. This increasing lack of first-hand communication of taxonomic knowledge is more than lamentable. Experienced knowledge and skills are desperately needed if biodiversity is to become known within a reasonable time frame. The severe shortage of human resources is acknowledged as a most

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serious constraint in reorienting systematics towards countering the biodiversity crisis (Boom, 1991; May, 1993). In 1991, as judged from the numbers of systematists teaching in UK universities, at least 60% of taxonomists were over 45 years old, with less than 10% under 35. The opposite situation typifies scientific disciplines such as physics with their majority of youthful practitioners (May, 1992; Krebs, 1992). Judged by on-going neglect, as exemplified by the current decline of systematics research and teaching in Canada (Idema, 1993) and South America (de la Penha, 1993), this situation is widespread, deteriorating and unresolved. The loss of expertise in systematics and taxonomy awaits recognition, let alone a pragmatic solution. The most serious result of insufficient recruits to the discipline is the loss of experienced knowledge possessed by established systematists, which is difficult to recover from serendipitous literature searches. Although a younger generation of systematists may confidently apply the powerful methodology of cladistics to systematics, all such research relies on expertise (invariably scarce and possessed by aging taxonomists) to identify organisms. Such knowledge cannot be assimilated with the requisite rapidity, nor easily recovered from the literature left by deceased experts. Direct exposure of students to the knowledge of experienced taxonomists is a cost-effective means of transferring this expertise. This experiential approach complements formal academic training, and is a highly efficient way of perpetuating and expanding expertise in systematics (Boom, 1991).

Decaying collections Biological collections have been termed libraries of life. In a climate of crisis, the values and paradoxical threats to natural history collections catalysed the formation of a World Council on Collections Resources (WCCR) which endorses: The central purposes of natural history collections are to record through specimens and related data the existence of species on earth along with their supporting geological structures, to carry out research on the interrelationships of plants, animals and minerals, and to communicate this knowledge to serve the needs of society... The preservation and conservation of natural history collections transcend local and national concerns. (Davis and Emery, 1993.) The overall mission and specific actions of the WCCR seeks to promote and co-ordinate the wise care, expansion and scientific use of natural history collections (Davis and Emery, 1993). It remains to be seen whether these actions will be implemented. The worldwide tragedy afflicting biological collections remains poorly recognized (Seymour, 1994). Only 0.5% of overall budgets of museums in the UK was spent on preserving natural history specimens in 1983 (Howie, 1986). The Natural History Museum in London spends less than 10% of its annual budget on preserving its 67 million specimens (Seymour, 1994). In 1985, a comprehensive survey revealed that at least two thirds of 254 museums in the UK (excluding Kew Gardens and the Natural History Museum) had lost parts of natural history collections through neglect, and that the integrity of seven million specimens was in jeopardy. Overall, museums were shown to be guilty of flagrant neglect of natural history collections, which had resulted in their wholesale decay (Williams, 1987). At least 2.5 billion natural history specimens exist worldwide. Preservation of many of these specimens is not ideal and most of those over 150 years old no longer exist. Considering the world's natural history collections as a whole, funding and support for even the most basic and essential requirements to preserve specimens is drastically


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inadequate (Howie, 1986, 1993; Hawkes, 1990; Seymour, 1994). A survey of mammal collections in North America revealed that even type specimens are inadequately preserved (Hawkes, 1990). This appalling neglect of specimens in natural history collections is most disturbing, primarily because they are vital to environmental conservation (NSB, 1989; Danks, 1991; Eldredge, 1992). At the very least, their loss will hamstring efforts to decipher the identities and origins of new specimens resulting from further exploration of the biosphere (Howie, 1993; Danks and Ball, 1993). While funding agencies dither and critics of the decline of systematics pontificate, neglected collections rot. Constituting the silent crisis in systematics, these losses of taxonomic skills and specimens proceed largely unrecognized. The extirpation of systematists' skills and scandalous neglect of collections has received insufficient attention within the numerous discussions of systematics' importance, despite the fundamental values identified in the science (Crowe et aL, 1989; Whitehead, 1990; Harvey, 1991; Eldredge, 1992; Krebs, 1992; Lane, 1992; Linnaean, 1992; Kim, 1993; Alberch, 1993; Janzen, 1993a, b). It is strange that two reviews (Krebs, 1992; Linnaean, 1992) of the state of systematics in the UK did not mention the comprehensive report on the country's collections (Williams, 1987). Inadvertently or otherwise, many systematists are guilty of ignoring the care of collections in favour of their use (Raven, 1988; Simmons, 1993). Natural history collections in many tropical countries appear at greatest risk, where neglect for their preservation and importance is symptomatic of more widespread lack of support for fundamental biological research, which: ... keep[s] institutions frozen in archaic methodologies because funding is never available when it is needed. The 'crazed ecology' philosophies pushed by the media and absorbed by the government have replaced effective work programs with bureaucratic 'paper ecology', creating what I call 'directionless environmental sciences'. Guessing has substituted hard science as a basis for most of the government's plans; soft talk has replaced knowledge; and a lack of scientific involvement in research and educational programs has created new myths and opened ground for erroneous concepts. (de la Penha, 1993.) The cost of collection, accession and curation of specimens has been estimated (Whitehead, 1990; D anks, 1991; Seymour, 1994). In particular, the future uses of botanical specimens have been questioned - particularly their future role in supporting identifications. The underlying motivation for this doubt of continued specimen preservation, especially those in large collections, is justifying maintenance costs to funding authorities (Waiters, 1993). The source of this anxiety over contemporary uses of collections is largely based in the unclear thinking and poor elucidation of the roles of preserved specimens in maintaining biological knowledge. The replacement of a reference book lost from a library is theoretically feasible - assuming at least one other copy of that edition exists. Conversely, the demise of even one historical specimen in a biological collection is irrevocable. Specimens are the material representation of unique information which can be interpreted by present and future biologists. The ravages wrought by the loss of specimens is a collective disaster which threatens the quintessential foundations of biological knowledge. Preserved specimens cannot be replaced. The millions of historical records in collections cannot be valued in conventional terms. Each specimen is historical, unique and inherently irreplaceable. The interrelated values of biodiversity and systematics and the latter's interrelationship to environmental conservation require elaboration to truly appreciate the serious consequences of neglecting the former. Most importantly, the role of taxonomic resources

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in maintaining biological knowledge requires explanation since the future of biodiversity depends on our knowledge of its complexity. We must examine in detail, how knowledge in a biological context is created and maintained.

Biological knowledge and the flow of information Knowledge and information are frequently synonymized, yet there is a definitive distinction between them. Their definitions hinge on a theory of knowledge articulated by Dretske (1981). The following discussion draws on Dretske's philosophy. Although some may construe this discourse as trite, it describes a far from trivial process but the pragmatic means of interpreting and managing information on biological organisms (complex objects whose categorization defies cursory observation) to build accurate knowledge. We need to acknowledge and rationalize the premise that biological knowledge relies on concepts based on empirical relationships between observed objects and facts. This premise is the prerequisite for quantifying origins and identities, and associations among organisms and other biological entities. Biological knowledge is built through the process of collating suites of related concepts. A concept is a belief based on aspects of information flow. Communication of this information, by written and electronic means, seeks to maximize accuracy and minimize equivocation. This information flows from perceived object to the human observer, who conceptually processes the received signals to interpret the natural entity as a concept. Information contained in a concept is specific to observer and object. Each concept possesses a specific etiology and individual concepts correspondingly differ in their etiological specificity. Concepts contain information of a high level of intentionality, structured as two facets each of which contains different information. One of these elements looks backward to informational origins; the other looks ahead to effects and consequences. No structure of belief qualifies as a concept unless it organizes information with these forward and backward aspects. As cognitive structures, concepts acquire their etiology from their informational origins, a content determined solely by the observed object's structure and origin - its informational heritage. In the case of a biological concept, such as the identity of an organism, information flows to the human observer from the particular organism of interest (preserved or living). A concept can be derived from such an organism at a particular point in time. Only if the organism is collected and preserved in a reference collection can other observers independently derive concepts of corresponding etiologies from this same object in the future. Observations reported on preserved specimens can be repeated and the resultant concepts independently checked. This facility is vital to quantify and compare their accuracy and equivocation. Preserved specimens are stable sources of reliable information (of minimal equivocation and maximum accuracy) from which biologists can assemble knowledge. Preserved specimens structure the concepts on which biological knowledge is built, so a cardinal requirement exists to maintain specimens to allow independent checking of concepts. In interpreting the complex natural world, preserved specimens unequivocally link together and focus the accuracy of information communicated by individual biologists to their peers. Humans are the singular, weak link in interpreting natural phenomena as causallyinduced concepts. For example, using aerial or satellite imagery to interpret spatial information and depict concepts as symbols on a map is susceptible to observer error, as are interpretations of the origins and associations of biological entities. In a similar way,


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visual observations of biological organisms can (inadvertently or otherwise) communicate unacceptable levels of noise and thus compromise accuracies of reported concepts (identities and origins of organisms). Concepts associated with specimens are inherently weakened if the latter are destroyed, to disappear from the domain of biological knowledge to erode our understanding of the natural world. Similarly, if the necessary means for their identification and classification are unavailable, derivation of new biological concepts from unrecognizable specimens obviously becomes problematic. In biology, the preservation of specimens in reference collections preserves knowledge. Cardinal requirements to account for the identities and classifications of organisms studied places a precedent on biologists to deposit voucher specimens in recognized biological collections (Davidson, 1952; Yates, 1985; Meester, 1990; Goldblatt et al., 1992; Marshall, 1993). The validity of many ecological, behavioural and physiological studies has been compromised because voucher specimens no longer exist or were never collected (Danks, 1991). Voucher specimens anchor knowledge, as they control the equivocation of all biological concepts. Their preservation permits concepts to be repeatedly checked, so published knowledge can be verified. The value of biodiversity Sustainable use, biological products and human perceptions The reliability and breadth of biological knowledge, ultimately dependent on taxonomic resources in its support of ecology and systematics, directly accounts for the values humans place on biodiversity. The values of biodiversity impinge on the human psyche, economics, health and agriculture (Wilson, 1992; Mittermeier and Bowles, 1993). Considerable value of biodiversity lies in its ability to counter adverse effects of contingencies such as unforeseen disease epidemics, and the vagaries of unpredictable climates and fluctuating economies (Burton et aL, 1992). It is how we come to value biodiversity that will decide the future of natural systems. Conferring minimal, or only aesthetic, value on terrestrial biodiversity will result in increasing economic demands for natural resources removing and replacing natural systems with the more productive and familiar landscapes we associate with agriculture and industry. 'The only certain way to save biodiversity is to use it sustainably.' (Janzen, 1992.) If biodiversity has value as a source of products, we will confer on it the status of valuable capital, and if maintained, this capital could supply a variety of valuable products to different markets (Wilson, 1992). Nevertheless, if perceived as valuable, such products must be tangible and of immediate benefit to society (Janzen, 1992). To utilize, and in this process save, biodiverse systems, we must understand their complexity. Increased knowledge of biodiversity, its accuracy dependent on taxonomic resources, will permit new and penetrating insights into the natural world and reveal a multitude of new opportunities to use biodiversity.

Sustainable utilization, conservation philosophy and knowledge of biodiversity A fundamental shift is occurring in our philosophy of environmental conservation, a change which concomitantly influences perspectives on biodiversity and biology (Botkin, 1991; Pickett et aL, 1992). Customary attempts to preserve areas of undefiled wilderness (and to maintain populations of conspicuous animals and plants within it) has been the prevailing conservation strategy, but particularly in the tropics is increasingly superseded by the active husbandry of conservation areas (Goodland, 1990; Janzen, 1992). For

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example, sustainable utilization of biodiversity in tropical countries seeks to involve human neighbours of conservation areas so they benefit from revenues derived from biological resources therein. Utilization of wildlife by local communities is becoming increasingly commonplace (McNeely, 1989). The philosophy of sustainable utilization is wholeheartedly embraced by Caring for the Earth; the Second World Conservation Strategy (IUCN, UNEP and WWF, 1991) and the Global Biodiversity Strategy (WRI, IUCN and UNEP, 1992), and forms the essence of the Rio treaties. McNeely and colleagues (1990) endorse the need for an increased knowledge of biodiversity, but are less specific as to the information required to aid its conservation. Over the past three decades, under the auspices of wildlife management, a considerable amount of knowledge has developed to manage natural systems, but is largely inadequate to support operations seeking to utilize biodiversity. The shift in conservation philosophy towards use of biodiversity now demands detailed knowledge of natural systems to locate, evaluate and sustainably use a variety of different organisms. Crossing this watershed in conservation philosophy, from preservation to utilization of biodiversity, places unprecedented demands on systematics. Scientific knowledge of many different species becomes essential to identify different organisms, understand their ecological and evolutionary relationships and summarize patterns of their occurrence. Taxonomic classifications universally communicate this detailed knowledge. From information supplied by ecology and systematics on relationships among organisms, investigators can accurately target certain populations (e.g. plants and their parasites) in searching for new biological products (Armbruster, 1992; Janzen, 1992, 1993a, b). This shift in information requirements is exemplified by the accurate and detailed knowledge required by biodiversity prospectors of a multitude of different species (Joyce, 1991; Reid et aL, 1993). It is very difficult to summarize accurately (because published information is comparatively inaccessible) our current knowledge of biological diversity. For example, taxonomists' efforts to describe new species over two centuries, as summarized by Hammond (1992) are a crude index of a gradual improvement in our knowledge of biodiversity at the population level. Knowledge of biodiversity at its higher hierarchial levels of organization (the structures of landscapes and functioning of ecosystems, sensu Noss, 1990) is principally improved by the ecological sciences (Soul6, 1990). The values of biodiversity, ecology and systematics are inextricably linked (Eldredge, 1992). They hold a synergistic value which can be conceptualized as humanity's knowledge of the multifarious complexities of biodiversity. In an inexorable socio-economic milieu, the values of biodiversity are invariably compared against the significance humans retain for the palpable benefits of industry and agriculture. To many minds, these benefits rarely extend beyond the amalgam of their immediate neighbourhoods. Forces, comprising economic processes and human motives and perceptions, will decide whether biodiversity is wisely utilized and by default, saved. Society's widespread ignorance, exemplified by the policy maker's and funding agency's poor perception of biodiversity issues, exacerbates the environmental crisis (Soul6, 1990; Ulfstrand, 1992; Myers, 1993). In all respects, scientific knowledge is essential to value and successfully manage the environment (Hardin, 1985). The power of systematics to classify and identify organisms and target products to be derived from them (Table 1) is all too often hidden from the observations of final consumers, whose perceptions invariably divorce any role of systematics in supporting agriculture and industry to produce biological products (Bryant, 1983). A multidisciplinary approach is needed to provide a broad


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scientific knowledge of biodiversity (Soul6, 1990; Wilson, 1992). Systematics and taxonomy constitute the dovetailed foundation to support such a mission. Their role in organizing and improving our knowledge of millions of interrelated organisms requires further examination.

The communication of biological knowledge

The scope of systematics Systematics is an essential science (Table 1). Its power to objectively interpret biological diversity and so understand the history of life on earth (Mayr, 1968) is increasingly recognized (Gould, 1986; Ridley, 1986; O'Hara, 1988; Lovejoy, 1993; May, 1993; de Queiroz and Gauthier, 1994). The overall goal of phylogenetic systematics is an accurate narrative of the evolutionary chronicle, by representing evolutionary events in their true context (O'Hara, 1988) and systematists are still engaged in finalizing the methodology and philosophy of their enquiry, as revealed by the lively debate, acrimonious rebuttals and acknowledgements of colleagues' philosophies and findings in journals such as Cladistics and Systematic Biology. Adopting the central concept of evolution in its philosophy, phylogenetic systematics, or cladistics, as the science of discovering biological history, aims to elucidate true relationships within clades of related taxa, by comparing patterns of inheritance of shared characters of organisms through time (O'Hara, 1988; Funk and Brooks, 1990). Phylogenetic information on the origins and evolutionary affinities of identified organisms confers considerable inferential and predictive power to biological investigations; knowing the phylogenies of related taxa is a prerequisite to understand biological diversity. Because they contain accurate information on evolutionary relationships, phylogenetic classifications form the basis for comparing interactions, associations and differences

Table 1. The values and interrelated services of systematics as an essential science in biology and industry Domain

Value and service

Identification

Taxonomies, as universal and accountable naming systems for organisms and biological taxa, are essential to organize and validate all biological knowledge

Information management Retrieval Comparison Prediction Theoretical biology Discovers Biological History

Existing knowledge on one or more taxa can be retrieved and compiled Properties of organisms and the communities and ecosystems which they comprise can be compared Existing knowledge of one or more taxa can be applied to infer and predict unknown properties of related taxa Narrates the evolutionary chronicle, by establishing the origins and relationships of life forms on earth and understanding how they have evolved (in response to changing environments)

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between organisms of different populations (Danks, 1988; Funk and Brooks, 1990; Wanntorp et al., 1990; Armbruster, 1992; Miles and Dunham, 1993). 'Comparative biology without a robust phylogeny is like chemistry without a Periodic Table.' (Crowe et aL, 1989.) The taxonomic method and verifiable biological knowledge The science of discovering, describing and classifying new biological taxa (in accordance with the Linnaean system of taxonomy, see below) constitutes the taxonomic method, and produces standard taxonomies. For over two centuries it has been the established process to communicate our knowledge of biological diversity. The methodology of taxonomy has been thoroughly reviewed by Mayr (1969), its protocols and philosophy corresponds closely to Dretske's (1981) theory of knowledge. Repeated examination of specimens (named according to a universal nomenclature) allows us to trace the origins and maintain the accuracies of concepts derived by taxonomists. The taxonomic method guarantees the veracity of biological facts in space, time and form to maintain and monitor an authentic knowledge of a diversity of life forms. Despite claims to the contrary (Hughes, 1992), it should also be noted that molecular and other techniques greatly complement but definitely do not replace the established taxonomic method, which designates and maintains specimens to account for biological knowledge (Goldblatt et aL, 1992; Krebs, 1992; Peterson and Lanyon, 1992; Banks et al., 1993). The cornplexities of biological information Biological information is incredibly complex since it pertains to the complexity of a huge variety of interrelated concepts (identities, classifications, and associated information) interpreted from organisms which interact within intricate systems in many different ways. Our current knowledge of biological diversity comprises numerous facts on over 1.4 million species. For over two centuries, this huge body of knowledge has accumulated in thousands of natural history collections and the biological literature. It comprises a vast number of parcels of interrelated information. This means that there '... is perhaps no other branch of science in which information storage and retrieval is as formidable and as crucially important a task as in taxonomy' (Mayr, 1969). Taxonomic databases Information comprising biological knowledge can be imagined as flowing from the environment, around the pivots (preserved specimens which structure unequivocal concepts) and into the exchangeable medium (in electronic format and the printed page) accessible to final users. Accuracies of these concepts can be independently checked by reexamining individual specimens, if they are still available in collections. Janzen (1993a) has traced the flow of such information, interpreted and managed by the taxonomic method, in a lucid example of how preserved specimens account for and maintain scientific knowledge of biodiversity. Systematics and taxonomy synthesize biological concepts (according to Dretske's theory of knowledge) to build universal frameworks of knowledge and any taxonomic classification is a storage and retrieval system for communicating biological information (Mayr, 1969; Mayr and Bock, 1994). The taxonomic method allows biological concepts to be compared, verified, synonomized, deleted and updated and taxonomies provide a robust framework within which biologists can channel accurate information from the environment to each other and society. Because taxonomic classifications link together a multitude of attributes on many


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different species, taxonomic databases should be available in an electronic format to be easily accessible to the user community (Janzen, 1992). An electronic database for a particular group of taxa can utilize a standard taxonomy, such as venomous snakes of the world (Golay et al., 1993) to organize the knowledge originally collected from reliably identified organisms preserved in collections. Information can be retrieved easily from such databases and packaged in a multitude of different ways to suit individual needs of a spectrum of users. Varied to consumer requirements, this knowledge may be collated into detailed distribution maps; scientific keys and illustrated field guides; origins and attributes of biochemicals and biomaterials for industry; and any body of facts on biodiversity as interpreted and marketed to society (Morin and Gomon, 1993). Once a taxonomy of a particular group becomes available, numerous avenues of opportunity are opened to society to interpret and use this knowledge (Janzen, 1993b). Unprecedented advances in computer technology radically lowers the costs of managing large quantities of complex information (ToNer, 1990), which has created considerable opportunities to generate and disseminate biological knowledge.

Identifications, taxonomies and the accessibility of biological knowledge Although discounted as a mere service, degrading the true value of systematics (Crowe et al., 1989; Renner and Ricklefs, 1994), correct identifications of organisms are the obvious, vital step towards investigating further aspects of the biology and relationships among taxa and the structure and functioning of natural systems. Correct identification is cardinal to all biology (Davidson, 1952; Mayr, 1968; Danks, 1988, 1991; May, 1993; Cotterill et al., 1994). Knowledge derived from identified specimens is the substance of further studies by palaeontologists, molecular biologists, systematists, ecologists and others (who operating under the collective umbrella of biogeography and evolutionary ecology) can elucidate why organisms occur where they do and how they interact (Eldredge, 1992; Danks and Ball, 1993). Traditional, or Linnaean taxonomy, when naming and classifying organisms, only estimates the evolutionary relationships among biological taxa and these classifications do not represent true phylogenies among the taxa they describe, as they are classified in a nested nomenclatural hierarchy. For this reason they are obviously unsuitable for phylogenetic comparisons (Miles and Dunham, 1993) and some authorities have suggested they are outdated. Ideally, a taxonomy should accurately communicate cladists' findings, but a universal system of phylogenetic taxonomy (communicating the evolutionary relationships elucidated by cladistics) has yet to be finalized. A phylogenetic taxonomy contains considerably more information about the taxa it represents, as its taxon names communicate evolutionary meanings, summarizing the origins and relationships of classified organisms (de Queiroz and Gauthier, 1994). Cladists will eventually classify, in their correct phylogenetic context, the taxa originally described and presently classified in Linnaean taxonomies. By placing the concept of evolution at the centre of taxonomy, phylogenetic classifications (of birds for example) are superseding standard taxonomies. Such replacements, although valuable and inevitable, must be carefully done because replacing a standard taxonomy with one based on cladistics can create difficulties in information retrieval (Mayr and Bock, 1994). As in any biological discipline, where investigators access and organize knowledge, cladists still rely on existing Linnaean classifications to obtain information and identify organisms. For example, phylogenetic estimation in the recent classification of venomous snakes of the world

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(Golay et al., 1993) contains inaccuracies. Nevertheless, the monograph organizes a huge quantity of knowledge to support investigations into all aspects of ophid biology, including the biotechnological potentials of venoms. Furthermore, this standard taxonomy is vital to future systematics research, aimed at an objective representation of phylogenies and a deeper understanding of snakes' origins and diversity. Despite their inadequacies, Linnaean taxonomies are the best available to communicate biological information (Mayr and Bock, 1994). A theory of knowledge based on information flow (Dretske, 1981), the circumstances of biological investigations (Mayr, 1968; Danks, 1988; see above), and the requirements for knowledge to conserve biodiversity (Raven and Wilson, 1992; Wilson, 1992; Lovejoy, 1993; May, 1993), places urgent requirements on Linnaean taxonomies, and the resources producing them, to universally communicate biological knowledge. The ultimate sources of this knowledge are the biological specimens collected during inventories of habitats.

Biological surveys and specimen collection The current emphasis on biological inventories has been dismissed as undermining the vitality of systematics (Renner and Ricklefs, 1994). Yet the deplorable state of our present knowledge of biodiversity, and the urgent need to rectify the situation, is well established (Raven, 1988; Wilson, 1992; Janzen, 1993a, b; May, 1993). Furthermore, the basic biology (phenology, behaviour, life history and habitat requirements) of the vast majority of described species is unknown, yet knowledge of their autecologies is essential to conservation (Lawton, 1993; May, 1993). Such information could be gathered during inventories (as in the BIOTROP methodology, Duellman, 1992), with research efforts co-ordinated to collect specimens and record ecological data on selected populations. Inventories are crucial to census the poorly known biosphere (e.g. NSB, 1989; NSF, 1992; Raven and Wilson, 1992; Wilson, 1992; Cotterill et al., 1994). They are a key process in improving biological knowledge, as the specimens they supply constitute fundamental information on biodiversity (Alberch, 1993). Inventories are '... not an end but rather a means to the end of efficient use of a repository of diverse objects too complex to be identified at a glance' (Janzen, 1992). Since they validate the concepts of knowledge, biological specimens reliably quantify biodiversity. With the possible exception of charismatic vertebrates (a trivial component of biodiversity), concepts of organisms' identities and species' distributions that are not directly referrable to preserved organisms (voucher specimens) cannot be substantiated. To succeed in knowing biodiversity, we need to appreciate the absolute necessity to collect living organisms and preserve them as specimens. This need requires clarification, as misunderstanding among some biologists (e.g. Hughes, 1992; Renner and Ricklefs, 1994) and muddied thinking within the conservation fraternity obscures the fundamental role of biodiversity inventories (and the taxonomic method) in supporting biodiversity conservation (de la Penha, 1993; Waiters, 1993). The deplorable lack of knowledge decrees actions, not sentimental objections to inventories (Cotterill et al., 1994). Many of the action plans addressing the biodiversity crisis entirely omit the need (Stuart et al., 1990) or do not sufficiently emphasize the fundamental role (McNeely et al., 1990; WRI, IUCN and UNEP, 1992) of taxonomic resources and systematics in biodiversity conservation. Requirements for inventories must be evaluated within the context of threats to entire habitats and ecosystems: so in effect,


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those who ignore this need denounce the cardinal roles of identification and classification in building biological knowledge. The future of natural history museums and herbaria

The unique attribute of natural history museums and herbaria are the collections they possess, a trait distinguishing them from other research institutions (Chalmers, 1992). Museums are places where collections are studied (Simmons, 1993). Taxonomy and systematics have traditionally been associated with these institutions. Nevertheless, Alberch (1993) asserts that museum operators should undertake more purposeful research and market themselves within the context of the biodiversity crisis. Natural history museums can supply a plethora of services (educational and other to society, Bodmer, 1990), but without natural history collections and their associated research, they lose the crux of their identity. If museums plan to become major contenders in biodiversity research, it may become increasingly difficult for them to continue their traditional vocation of offering a wide range of services, ranking each of equal importance. It must be realized, perhaps enforced, that museum activities have to be prioritized. As an overall priority, those accountable for these institutions must support the preservation of collections. It remains to be established whether donors will fund systematics in institutions which attempt to service a variety of society's whims, at the expense of taxonomic resources. Society's attitude towards natural history collections and associated research can be gauged by the fact that the funding available to preserve natural history collections was less than 10% of that allocated to the UK's art collections in 1983 (Howie, 1986). Similarly, research (estimated by numbers of publications on matters of biological collections' management and preservation) is exceeded ten-fold by related research into art collections (Howie, 1993). If we are to be honest with ourselves and face reality, we must evaluate taxonomic resources against the global scale and singular austerity of an environmental crisis: the value of everything else that museums stand for pales into insignificance compared with taxonomic resources. We patently cannot afford to allow taxonomic resources, as the foundations of biological knowledge, to decline any further. Their international value reaches way beyond the myopic and parochial policies typical of many herbaria and museums, which invariably sway to the tune of national politics and in groping for survival, create or adapt corporate policies as an answer to climates of dwindling support. Corresponding to the environmental crisis, there is an urgency to actively market systematics and the value of taxonomic resources to society (Chalmers, 1993; Emery, 1993; Hoffmann, 1993; Lovejoy, 1993; May, 1993). A healthy state of taxonomic resources is crucial to such a campaign and industry, where collections (and the skills to interpret them) will realize increasingly global values. Museums which cling to national strictures will become even more vulnerable if they ignore international demands for biological knowledge, the needs for which at least equal that of physics and medicine, which routinely receive considerable funding. Restructure or die? Many of the corporations, which represent museums and herbaria, require reorganization and expansion if research on biological diversity is to become their central raison d'Otre.

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Supported by professional management, radical increases in funding and manpower are necessary to support such goal-orientated research into biodiversity. In many institutions, administrations must be restructured if taxonomic resources are to be properly supported and research is to be encouraged. In orientating activities towards researching biodiversity at a global scale, many museums will need to replace antiquated bureaucracies, inherited from the 'smoke stack era of the last century', with the novel and more flexible organizational structures described by Toffler (1990) as typical of corporations operating efficiently in the communication age. Operators in museums and herbaria must be able to communicate effectively in the international theatre and be capable of making responsible decisions in their operations. Taxonomists and systematists typically study their organisms of interest across the entire spatial range of their distributions, at continental and global scales, and in consequence - because organisms' distributions are rarely constrained by political boundaries - this research is typically international in its scope. With objectives to understand tropical biodiversity and directly facilitate its sustainable use, biodiversity research centres such as InBio, Costa Rica's National Biodiversity Institute (Gamez et al., 1993), are recent departures from the entrenched traditions of most museums. The InBio experience, in supporting goal-orientated systematics research, aimed at the marketing of biodiversity (Janzen, 1992; Gamez et al., 1993; Reid et al., 1993) demonstrates that taxonomic resources can be effectively deployed outside the traditional domain of museums and herbaria. An increasing trend in North America adapts corporate structures of natural history museums to support the sound management of collections, and so preserve the core of these institution's identities (Simmons, 1993). As they stand, most museums and herbaria lack the requisite reserves of initiative, flexibility and efficiency needed to practise systematics with the fervour dictated by a global environmental crisis. Institutions that fail to reorganize themselves adequately may flounder, but their taxonomic resources are too valuable to disregard. A less selfish solution for recalcitrant institutions would best place neglected taxonomic resources within more mission orientated institutions, similar to Costa Rica's InBio.

Why is systematics so neglected? The megafaunal bias The bulk of species comprising biodiversity (invertebrates and the microbial fauna) receive insufficient attention from the conservation fraternity (Kellert, 1993; Samways, 1993). Neglect of small bodied organisms is an integral property of the biodiversity crisis, and has been labelled the 'megafaunal bias' (Platnick, 1991). It is typified by the public's feverish devotion to whales, pandas, elephants and rhinos. The megafaunal bias characterizes conservation activities in Africa, where according to popular opinion, large mammals apparently exist in serene splendour amidst pristine wilderness (Adams and McShane, 1992). Blinkered in their subjective focus and frequently masquerading under the umbrella of biodiversity conservation, well-intentioned donors expend considerable resources on studying and trying to save the obvious or cuddly vertebrates, often to the detriment of biodiversity conservation. A serious dearth of information on invertebrates in a review of the status of Afrotropical biodiversity (Stuart et aL, 1990) typifies this sentimental subjectivity. The focus on charismatic animals ignores the complex issues (socio-economic and biological) involved in biodiversity conservation and the more encompassing and vital requirement to save and thoroughly know habitats and ecosystems


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(Janzen, 1987; Soul6, 1990). A widespread failure to acknowledge the complexities and true scope of biodiversity (Kellert, 1993), may explain society's poor appreciation of, and the consequent neglect of taxonomic resources. This is a considerable obstacle to the objective study of biodiversity.

Inadequacies of biological curricula The uses of biological specimens extend beyond their central role in validating organisms' identities and origins. Biological specimens have supplied data to countless quests in ecology, behaviour, forensics, pollution studies, industry and agriculture (Bryant, 1983; Danks, 1988, 1991; Krebs, 1992). Yet, the majority of biologists working outside systematics appear unaware of such values of natural history collections. Many biologists ignore the fundamental role of voucher specimens in the scientific investigation of the natural world (McAlpine, 1986). Neglect of this fundamental requirement has compromised the scientific validity of numerous investigations (Meester, 1990; Goldblatt et al., 1992). Lack of cognizance of these issues is a serious indictment on the many biologists and environmentalists who disregard the relationship of taxonomic resources to verifiable biological knowledge. The neodarwinian paradigm in evolutionary biology, through its focus on molecular biology and genetics, has foregone efforts to understand biological complexity at the organismal level (Margulis, 1990). This bias possibly reflects on systematics' poor marketing of itself and the contents of biological curricula in high schools and universities. Gould (1986, 1989) and O'Hara (1988) have argued for a greater cognizance of history by biologists, and for its thorough integration with biology. Although popular opinion may suggest otherwise, the comparative method in evolutionary biology is not inferior to the orthodox experimental method in the physical sciences, and in fact, both investigative approaches borrow each other's techniques and methodologies to quantify historical events in evolution. Biologists cannot ignore history's role; the way unique evolutionary events produced the living organisms comprising natural systems. Biological curricula need to articulate the value of studying life's history, to emphasize that by studying extinct and extant organisms and environmental change - through elucidating the timings and characteristics of biological events - systematics discovers this history and narrates the evolutionary chronicle. Above all, the singular value of taxonomic resources as the foundations of biological knowledge must become more widely known and respected. Salvage of taxonomic resources is critical

When we think of systematics and significant biological collections, the major museums of Europe and North America hold our attention. These are undoubtedly important, but many collections exist in other countries, including the tropics, and not all of them are in museums. The integrity of many such collections (particularly those under government 'care') are in serious jeopardy (de la Penha, 1993). In several African countries, for example, numerous private collections of insects, birds' eggs and plants are nurtured by zealous owners. Essentially anonymous, many such repositories have grown through a process of inter-generational devotion and inheritance, and bequeathed collections of international scope and importance. As each of their specimens is unique, they hold an importance equal to those collections in museums. Accessible, or untapped, the knowledge upheld by all these sources is indispensable to study biodiversity.

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Research by taxonomists and systematists has been too fragmented, lacking an essential synergism (Alberch, 1993), such that we do not even know exactly how many or what species have been described. A major reason for this uncertainty is that the necessary information, especially at global scales, is not sufficiently accessible (May, 1992; Barkley, 1993). Existing published information and collections and improvements in biological knowledge (inventories and allied research) needs to be structured into a framework of goal directed activities (Table 2). To marshall existing resources, this mission will obviously depend on the requisite technology and sufficient funds. Collaboration with expertise in other scientific disciplines and industries is vital to extract and propel the information, encoded in biological specimens, to users of the environment. Pluralistic research is called for. Efficient dissemination and exchange of this knowledge is as vital. So such efforts, linked at a global scale by using the rich potentials of information technology and global communications networks, are critical to process biodiversity information efficiently. A Biodiversity Information Network (see Canhos et al., 1992) should greatly improve accessibility to biological knowledge. Formation of the 'taxasphere' (reliant on information technology and in place to remove 'taxonomic roadblocks') is envisaged by Janzen (1993a) as critical to explore the complex biodiversity of a poorly known biosphere. Before an efficient taxasphere can be formed, let alone begin to function, the silent crisis destroying taxonomic resources must be recognized and halted. The expanding dialogue on the importance and plight of systematics and especially the fundamental requirements to improve and sustain biodiversity knowledge decrees serious consideration and urgent action on a sequence of nested activities (Table 2). Conclusion: first step before a giant leap Taxonomic resources support each and every biological discipline and maintain the integrity of all biological knowledge, their benefits extending tentacle-like into every biological activity (Mayr, 1968). The development of cladistics, founded by Hennig (1966) and allied with unprecedented advances in computer technology and molecular biology over the past three decades, is singularly responsible for considerable opportunities and growth in phylogenetic systematics (Doyle, 1993). Since the concepts sustaining our knowledge of life's complexity rely on specimens to validate narratives of evolution, systematists' investigations will increasingly rely on natural history collections, particularly since new techniques allow direct investigation of their preserved nucleic acids; a development which permits a whole suite of questions on phylogenetic relationships to be investigated (Krebs, 1992; Doyle, 1993). Wilson (1989) has called for the stewardship of systematics within the biological sciences in the pluralistic exploration of diversity. Such pluralistic research, reliant on systematics, will obviously fail if the foundations (collections and the expertise to interpret them) are lost. Interest and concern over the high profile subject, biodiversity, is no longer limited merely to biologists. It is less than pitiful that the scientific procedures of studying and building biological knowledge are so badly understood and poorly supported, not only by society, but by the majority of biologists. Widespread disregard and ignorance of biology's pivotal dependency on taxonomic resources is more than tragic. Exploration of diversity is the domain of future opportunity in biology (NSB, 1989; Wilson, 1989, 1992; Law et al., 1992) and the outcome of our efforts to maintain the integrity of the biosphere ultimately depends on successful organization and expansion of our knowledge of biodiversity. The

Action Assess number, origins and condition of specimens Questionnaires, and literature reviews to enumerate taxonomists and collections managers Promote and facilitate international collaboration and research a. Improve biological curricula b. Experiential training of graduates by established experts a. International funding of taxonomic resources and research b. Trust funds established to guarantee sustainability of institutions housing taxonomic resources c. International monitoring of the administration and performance of corporations supporting taxonomic research

Objectives

1. Assess biological collections

2. Assess taxonomic expertise

3. Optimize research efforts

4. Increase quality and quantity of skills to study and maintain collections

5. Avail sufficient funds for continued maintenance of collections

a. Funds b. International legal agreements, collaboration and enforcement of agreements

a. Funds b. Sanctioning of actions by authorities responsible for institutions

a. Funding for travel and communication b. International agreement and national support

Resources and expertise to collate information

Taxonomic experts, collection managers, questionnaires and international collaboration

Resources and conditions

Integrity of taxonomic resources and biological knowledge guaranteed

Sufficient expertise to study and manage collections

Productive research into biodiversity at continental and global scales

Lists of experts and deficiencies in expertise identified

Overview of status and condition of collections

Outputs

Table 2. Actions and resources required to assess taxonomic resources, salvage biological collections, and sustain biological knowledge

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accuracy and reliability of this knowledge ultimately depends on the integrity of specimens in biological collections, and the abilities of biologists to interpret their identities and affinities. It needs to become generally known that biological knowledge is built of individual concepts, whose accuracy is anchored by preserved specimens - the philosophy of this argument being based on a theory of information flow (Dretske, 1981). Since the maintenance, improvement and application of this knowledge depends on the integrity of natural history collections, the availability of human skills to maintain and study the specimens therein is critical. Threats to collections and the serious decline in these skills will severely stifle efforts to sustain and build a coherent body of knowledge of biodiversity. The Convention on Biological Diversity (UNEP, 1992) prescribes that its signatories know the status of the biodiversity in their countries in order to support the in situ conservation of habitats and ecosystems. This, and additional exactments on taxonomy (NSB, 1989; Linnaean, 1992; NSF, 1992; Wilson, 1992; Alberch, 1993; Danks and Ball, 1993; Davis and Emery, 1993) are paradoxical in view of the discipline's disintegration. Janzen (1993a, b) has argued that taxonomy and systematics should be supported as central activities in a nation's policy of environmental conservation, because they are crucial to maintain and manage tropical biodiversity. Recent assessments of research priorities for biology place a precedent on supplying information for environmental decision making. The Diversitas programme of UNESCO (Robertson Vernhes and Younes, 1993) and the Sustainable Biosphere Initiatives (Sala, 1992) categorize an integrated suite of questions into action plans for ecological research. In North America, there are welcome signs of a revival of systematics allied to these international research agendas in ecology. Raven and Wilson (1992) have outlined the essential infrastructure and conditions necessary for a fifty-year survey of global biodiversity. Formative plans are laid for an 'All Taxa Biological Inventory'; a thorough study of the biodiversity within a selected neotropical site (Yoon, 1993). A global mission statement - Systematics Agenda 2000 - was recently announced (Anonymous, 1994; Mcilwain, 1994) which aims to marry the unrealized scope of the investigative and predictive powers of systematics and taxonomy with the needs of society and so realize an interrelated set of goals which the discipline has always lacked (Cracraft and Hoagland, 1991). It cannot be overemphasized that correct knowledge is vital to manage and protect biodiversity and avoid unpleasant repercussions, where far-reaching decisions on environmental issues are based on false inferences and unsubstantiated observations. It is seldom acknowledged that environmental decision-making places a tremendously heavy responsibility on us. We need accurate, precise information to decide how much and what biodiversity we bequeath to tomorrow's generations. A greater synergy between environmental management and biology is needed; to tightly weld systematics and taxonomy into the forefront of issues in biodiversity conservation; discovering, naming and classifying organisms and the places on earth where they occur. To be maintained, the landscapes and ecosystems which these organisms comprise must become better known, appreciated and valued. Should we succeed in halting the biodiversity crisis, humanity will achieve a truly mature relationship with the biosphere. Rescuing and consolidating taxonomic resources will provide for the future of biological knowledge and lay the foundations on which to launch this herculean task.


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Acknowledgements R.D. Barnes, N.C. Bennett, D.G. Broadley, A.N. Channing, T. Chatwin, D.N. Cotterill, J.M. Dangerfield, R.C. Drewes, A.J. Gardiner, M. Gardiner, J.R. Ginsberg, D.L. Harrison, R. Hoffman, C.W. Hustler, M. Kruger, R.D. McDairmaid, J.L. Minshull, E.M.B. Parry, S. Schilling, A.L. Sparrow, B. Stergios, and M.H. Wilkens kindly commented on drafts of this paper and made valuable suggestions. I am grateful to Alan Channing for emphasizing the differences between the phylogenetic and Linnaean systems of taxonomy. Any errors and misinterpretations are my own.

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