10841_2006_9018_11_1-web 47..59 Abstract Studies of conservation biology involving tiger beetles have become increasingly common in the last 15 years. Governments and NGOs in several countries have considered tiger beetles in making policy decisions of national conservation efforts and have found tiger beetles useful organisms for arguing broad conservation issues. We trace the evolution of the relationship between tiger beetle studies and con- servation biology and propose that this history may in itself provide a model for anticipating developments and improvements in the ability of conservation biol- ogy to find effective goals, gather appropriate data, and better communicate generalizations to non-scientific decision makers, the public, and other scientists. According to the General Continuum of Scientific Perspectives on Nature model, earliest biological studies begin with natural history and concentrate on observations in the field and specimen collecting, fol- lowed by observing and measuring in the field, manipulations in the field, observations and manipu- lations in the laboratory, and finally enter theoretical science including systems analysis and mathematical models. Using a balance of historical and analytical approaches, we tested the model using scientific studies of tiger beetles (Coleoptera: Cicindelidae) and the field of conservation biology. Conservation biology and tiger beetle studies follow the historical model, but the results for conservation biology also suggest a more complex model of simultaneous parallel developments. We use these results to anticipate ways to better meet goals in conservation biology, such as actively involv- ing amateurs, avoiding exclusion of the public, and improving language and style in scientific communi- cation. Keywords Cicindelidae Æ Conservation biology Æ History Æ Models Æ Tiger beetles Introduction The early 20th Century Spanish philosopher, George Santayana, is credited with the quotation, ‘‘Those who cannot learn from history are doomed to repeat it.’’ Although easily dismissed as a trivial aphorism, is it possible that this statement constitutes a testable hypothesis that we can use to understand and antici- pate advances in sciences such as conservation biol- ogy? Conservation biology is a field with too few years of experience to have engendered broad interest in its past (Zirnstein 1996; Meine 1999; Siemann 2003). However, its history together with that for longer- established supporting fields, such as systematics, genetics, wildlife management, and ecology, may hold critical information for developing future directions and goals for conservation biology. Faced with con- stant shortages of funding to adequately gather infor- mation and conduct studies, lessons from history may be useful as another set of tools in the quest for meeting these goals (Maienschein 2000; Gaddis 2004). CXLV, Studies of Tiger Beetles D. L. Pearson (&) School of Life Sciences, Arizona State University, Tempe, AZ 85287-4501, USA e-mail: dpearson@asu.edu F. Cassola Via F. Tomassucci 12/20, I-00144 Rome, Italy J Insect Conserv (2007) 11:47–59 DOI 10.1007/s10841-006-9018-9 123 B E E T L E C O N S E R V A T I O N Are we doomed to repeat history? A model of the past using tiger beetles (Coleoptera: Cicindelidae) and conservation biology to anticipate the future David L. Pearson Æ Fabio Cassola Received: 23 September 2005 / Accepted: 20 December 2005 / Published online: 7 November 2006 � Springer Science+Business Media B.V. 2006 In a search for patterns within the history of scientific studies, historians have analyzed several fields from physics (Nye 1996) to biology (Killingsworth and Palmer 1992). Are there steps common to all scientific en- deavor? What recognizable patterns of change take place, and what are the significant factors causing the changes? How can they best be compared? Apart from satisfying intellectual curiosity, a solid understanding of patterns in the development of science could prove useful for conservation biology in many ways. It could: (1) help determine priorities for funding agencies, (2) enable biologists to better communicate with and inform non-scientific decision makers, (3) focus individual re- searcher goals, (4) prepare cooperative research agen- das, (5) formulate more reliable and efficient models for management and conservation goals, and (6) help anticipate problems that can then be ameliorated. Methods The historical model History does not lend itself to experimental repeat- ability (Gould 1989), and thus tests of patterns in his- tory rely on alternative methods. One of the most reliable techniques for answering pertinent historical questions and testing for patterns is by using insights from one field to tell us something about another—a process called consilience by historians. In so doing, we can make sense of the past and perhaps anticipate the future (Gaddis 2004). Within biology, such patterns have been proposed for understanding the historical progression of human cultures. Important causes with consistent outcomes across unrelated cultures include environmental factors (Rolett and Diamond 2004), plant and animal domestication (Diamond 2002), dis- ease (Acemoglu et al. 2001) and food production (Hibbs and Olsson 2004). Along the same lines, one general model of the his- tory of science proposed to anticipate historical pat- terns in biology is the General Continuum of Scientific Perspectives on Nature (GCSPN) (Killingsworth and Palmer 1992). According to the GCSPN, earliest bio- logical studies begin with natural history and concen- trate on observations in the field and specimen collecting, followed by observing and measuring in the field, manipulations in the field, observations and manipulations in the laboratory, and finally enter the- oretical science, including systems analysis and mathe- matical models. What is not clear is whether each of these chapters in the development of science can be identified as a chronological step or phase, or even more controversial, whether each step has identifiable and quantifiable characters that can be used to estimate the maturity of the field of study (Farber 2000). In addition to these uncertainties, this model has other constraints. As with all models, simplification is an acceptable aspect of their use as long as the results are interpreted within these limitations. Also, similar to many ecological and landscape studies, using time intervals that are too small or too large can obscure important patterns. Finally, sociological, economic, and psychological forces can be more crucial in affecting models of temporal changes than generally realized, but these factors often are difficult to incor- porate into general models. With these assumptions in mind, Battalio (1998) listed a series of specific char- acters that would demonstrate historical steps within the GCSPN model: (STEP 1) Descriptive natural history and search for new species predominate (STEP 2) Now an experimental science rather than a natural history model (STEP 3) Power is transferred from expert amateurs to trained professional scientists, and graduate training for employment in the field has become available (STEP 4) Systematics no longer dominant, and re- search focused more on theoretically complex issues with extensive use of graphs and statistical inference in publications (STEP 5) Formation of research teams and increasing evidence of socialization, such as use of acknowledg- ments sections, associations of peers, and co-authored publications (STEP 6) Technical terminology and methodology so refined they now limit the audience that can fully comprehend them (Fig. 1). Test subjects The history of entomology provides a rich and varied set of potential subjects to test the model. However, Fig. 1 Linear progression of steps in GCSPN model in which each step replaces the former one 48 J Insect Conserv (2007) 11:47–59 123 because of the myriad and often independent histories of various insects groups within entomology (Sorensen 1995), we felt that an initial test of the GCSPN model would be more manageable by using a single group. Tiger beetles (Coleoptera: Cicindelidae) provide a relatively discrete taxonomic unit whose history is well documented (Pearson and Cassola 2005). The tiger beetles are a small but distinct group of over 2600 species whose biology is also well known (Pearson and Vogler 2001). These beetles are attractive, fast-flying and fast-running insect predators that occur in many diverse habitats around the world. Many of the same characteristics of tiger beetles that have generated considerable interest among amateurs and professional biologists have also contributed to their increasing role in conservation studies. Most important among these characteristics is the ease with which most species can be found and identified in the field, their habitat specificity, and their value as indicators of habitat health and of biodiversity. Also, because they have been well-collected and studied, their past and present distributions are known sufficiently to evaluate historic trends of decline in range or abundance (Desender et al. 1994; Knisley and Fenster 2005). In addition, we will use the history of the field of conservation biology as a test subject. With a combi- nation of narrative and comparative analysis, we pro- pose to compare these two histories to test the validity of the model. Finally, we pursue the possibility that if the resultant pattern of steps conforms to the GCSPN model, can the model and its assumptions be used to anticipate and direct future steps in conservation biology? Results Step 1: Descriptive natural history and search for new species predominate As claimed by the GCSPN, much of the earliest history of conservation biology revolved around documenta- tion of species, in this case their extinctions. In the late 18th Century, American authors Ralph Waldo Emer- son and Henry David Thoreau influenced the devel- opment of Transcendentalism, a philosophy associated with nature. Through their writings, preservation of nature and wilderness became a powerful, novel doc- trine. In the midst of manifest destiny and impressions of inexhaustible resources, the unexpected disappear- ance of once abundant species, such as the Passenger Pigeon, and near extinction of the American Bison, first made extinction seem a real possibility, and the causes of extinction of individual species became an important area of study for the nascent field of con- servation biology. Because of an extensive knowledge of taxonomy and distribution starting with Linné (1758), tiger bee- tles lent themselves to early studies of declining pop- ulations and extinctions. As such, several species and populations of tiger beetles became some of the first insects declared legally endangered or threatened with extinction. Pearson et al. (2005) estimate that at least 33 (15%) of the 223 named species and subspecies of tiger bee- tles in Canada and the United States may be declining at a rate that justifies their consideration for inclusion on the US Fish and Wildlife Service’s List of Endan- gered and Threatened species (Fig. 2). However, at present, only four of these are officially listed by the federal government, and several others are under consideration for listing. In addition, several other countries (Belgium, Canada, Germany, Great Britain, Lithuania, The Netherlands, South Africa and Sweden), at least 24 individual states and provinces within the United States and Canada, and international NGOs (World Conservation Monitoring Centre and IUCN) have developed lists of endangered and threa- tened species that include tiger beetles. Few insects are well-enough known globally to document these types of population decline. Because of the rich collections of tiger beetle specimens avail- able for study, however, the disappearance of species from former parts of the range can be authenticated. From these historical records, some long-term changes in the environment can also be deduced (Nagano 1980; Desender and Turin 1989; Desender et al. 1994; Yarbrough and Knisley 1994; Kamoun 1996; Trautner 1996; Berglind et al. 1997; Diogo et al. 1999; Knisley and Hill 2001; Richoux 2001; Sikes 2002; Goldstein and Desalle 2003; Horgan and Chávez 2004; Mawdsley 2005). Thus, tiger beetles help offer a window into our past and can provide insight as to where protective measures are needed (Babione 2003). Step 2: Now an experimental science rather than a natural history model For tiger beetle studies, the major intellectual advance during the last half of the 18th Century was an often- conflicting attempt to place the growing number of species into a natural array of groupings. By moving from pure description to evolutionary questions, these attempts at phylogenetics were also some of the first signs of a change into an experimental paradigm (Barrow 1998). With more species known, better J Insect Conserv (2007) 11:47–59 49 123 chances for comparisons, and greater competition for research subjects among the increasing number of experts, tiger beetle systematists ventured into more sophisticated areas of research. Field naturalists such as A.R. Wallace and H. W. Bates often collected tiger beetles wherever they traveled. Emergent but signifi- cant ideas about behavior, ecology and evolution also grew from their experiences of collecting and observ- ing these beetles. The German medical doctor, Wal- ther Horn, became the greatest authority and acknowledged specialist of the tiger beetle family, working almost solitarily for more than 50 years. Although predominantly taxonomic in nature, his articles began, later in the 1900s, to incorporate experimentally testable ideas of habitat, biogeography and intraspecific variation (subspecies). Besides reconstructing the past, tiger beetles are useful for conservation in other ways. Because of political, sociological and economic pressures, conser- vation policy and research are under pressure to pro- duce quick results. This pressure is so pervasive, and the time, money and personnel to do the work are so limited that conservation biology is called a ‘‘crisis discipline,‘‘ in which risk analysis has become a major element (Maguire 1991). A common approach to resolving these problems has been to use indicator taxa as test organisms that purportedly represent other taxa in a complex environment. By focusing studies on a small but representative subset of the habitat or eco- system, patterns of habitat degradation and population losses can be more quickly and clearly distinguished (Noss 1990). Unfortunately most taxa suggested for use as indi- cators have been selected primarily on the basis of their public appeal (Pearson 1994). The consequences have cast doubt on the general usefulness and accuracy of bioindicators in conservation policy-making. For instance, among animal taxa, most studies using indi- cator taxa have relied on vertebrates, especially those ‘‘species of high public interest’’ (USDI 1980). Verte- brates, however, tend to be relatively long-lived, have low rates of population increase, long generation times, and comparatively low habitat specificity (Murphy et al. 1990), all of which tax the time and finances for proper investigation. As a result, there is a trend now to rely more and more on arthropod species, especially insects, instead of, or in addition to, vertebrates as appropriate indicator taxa (Pyle et al. 1981; Kremen 1992; Samways 1994; McGeoch 1998). Tiger beetles have been used throughout the world to test and develop better guidelines for choosing bioindicators (Holeski and Graves 1978; Schultz 1988; Bauer 1991; Pearson and Cassola 1992; Rivers-Moore and Samways 1996; Kitching 1996; Rodrı́guez et al. 1998; Cassola and Pearson 2000; Cassola 2002; Arndt et al. 2005). First, the category of bioindicator is determined (Kremen et al. 1993). Will it be used for monitoring (Greenberg and McGrane 1996), in Fig. 2 Controlled area in Santa Cruz Co., California, to protect the officially endangered Ohlone Tiger Beetle (Cicindela ohlone) Photo courtesy Univ. Calif. Santa Cruz Grounds Dept. 50 J Insect Conserv (2007) 11:47–59 123 inventory (Lees et al. 1999), as an umbrella (Mitter- meier et al. 2004), or some other type of model organism? Then a claim is made that a species or taxon, such as tiger beetles, is ideal as a bioindicator in a specific category. That leads to tests of whether this proposed indicator taxon meets the demands of widely accepted logistical and biological criteria for ideal indicators within each category. A useful bioindicator taxon should have characteristics such as stable tax- onomy, well-known biology and readily observed and manipulated (Brown 1991). More recently, it has be- come evident that even when chosen carefully, a single taxon is unlikely to be adequate. Seldom will a single taxon reflect accurately an entire habitat or ecosystem (Ricketts et al. 1999). Choosing a suite of indicator taxa from different trophic levels or different subhab- itats within the area of interest probably produces better data on which to base rational and informed biological and policy decisions. Nevertheless, each of the suite of candidates should be vetted experimentally to determine its appropriateness for that specific use as a bioindicator. Step 3: Power is transferred from expert amateurs to trained professional scientists, and graduate training for employment in the field has become available In the late 1800s, the first conservation organizations, such as the Audubon Society and the Royal Society for the Protection of Birds, were formed with both pro- fessional and amateur participants. In the next few decades, the work of these professionals and amateurs created many conflicts, such as the benefits of specimen collecting and use of common names. Little by little, professional academicians and government employees with advanced degrees, such as Aldo Leopold and Rachel Carson, took over the study and communica- tion of conservation problems. In 1985 the Society for Conservation Biology was established, and by 2000 it had 5100 professional members. Conservation studies that involve insects have become more common (Bossart and Carlton 2002) in recent years, often focused by international insect organizations, such as the Xerces Society for the Conservation of Inverte- brates. In 1997 the Journal of Insect Conservation was launched in conjunction with the British Butterfly Conservation Society. By this time additional national societies dedicated to the conservation of insects had been formed in Asia, Europe and North America. Along with journals focused on this area, graduate programs and salaried positions as conservation biol- ogists, many of whom use insects as test organisms, became established, and the leadership and predomi- nance of professionals became more and more obvious. For tiger beetles, the near monopoly of a single expert, Walther Horn, had great influence on the direction of studies. Beyond his tight control of tiger beetle taxonomy, however, a few other professional biologists began to publish scientific articles using tiger beetles as test organisms for geological history (Wickham 1904), ecology (Shelford 1907), and behav- ior (Shelford 1902). The use of tiger beetles in con- servation did not begin until the late 20th Century (Pearson and Vogler 2001), and some potentially divisive problems, such as the development of common names, were less disruptive among amateur and pro- fessional tiger beetle workers (Pearson 2004) than with other groups, such as birds, butterflies and dragonflies. Even more subtly, professionalization of scientific articles, including those for conservation biology and tiger beetle studies, is reflected in its evolving language, writing styles, and grammar. Linguistic analysis of journals and scientific articles shows consistent changes that indicate levels of expertise and establish levels of authority, further separating professionals from ama- teurs. Some examples of changing words include adverbs that show degrees of reliability, such as ‘‘undoubtedly’’ and ‘‘possibly,’’ induction, such as ‘‘must’’ and ‘‘evidently,’’ identification of hearsay evi- dence, such as ‘‘it seems’’ and ‘‘apparently,’’ reserva- tions of deduction, such as ‘‘presumably’’ and ‘‘could,’’ and hedges, such as ‘‘approximately’’ (Chafe 1986). In addition, professional science writers use distinctive writing devices that include reduced use of personal pronouns, reliance on passive voice, a decrease in the number of simple sentences, the presence of technical terminology, an emphasis on reliability of evidence, and the use of citations (Lakoff and Johnson 1980). Carter (1990) also showed that although professionals rewriting scientific articles for semi-popular or popular consumption tend to write in broader generalities and use methods more similar to amateurs, they retain a concept of domain-specific knowledge that distin- guishes them from the style of amateurs. Step 4: Systematics no longer dominant, and research focused more on theoretically complex issues with extensive use of graphs and statistical inference in publications Among tiger beetles, in areas other than taxonomy, the 1960s saw a relatively small increase in articles published on behavior, ecology, morphology, bioge- ography and ecology (Pearson 1988). But starting in the 1980s, physiological studies of tiger beetles J Insect Conserv (2007) 11:47–59 51 123 emerged (Dreisig 1980; Hadley et al. 1988; Gilbert 1997; Hoback et al. 2000; Okamura and Toh 2004). In the 1990s, genetics studies began to appear (Galián et al. 1990; Proença et al. 2002), and by this time, these and other non-taxonomic publications constituted 85% of the articles on tiger beetles with statistical proce- dures and graphs. One area in which tiger beetles were at the forefront of more complex conservation biology studies was in the statistical application of assumptions of depen- dence among data points. In initial comparisons of species patterns across regions and countries, Pearson and Cassola (1992) claimed that among the tested attributes of tiger beetles as an ideal bioindicator was a high correlation between their species numbers and those of other groups. If one goal is to establish con- servation areas with the highest species diversity, tiger beetles were very useful because where you found more of them you also found more of other species like birds and butterflies. But tiger beetles, at the right season, could often be surveyed in a few weeks whereas birds took years to survey adequately in the same area. In addition, it was easy to train students and local workers to observe and sample tiger beetles, but training these same people to observe other taxa, such as birds and butterflies, was an enormous undertaking. Thus, one could argue that tiger beetles are logistically useful and biologically appropriate candidates to help represent entire habitats or ecosystems for species inventories. A major problem, however, was the misapplication of a common simplifying component in statistical tests used by many biologists (Carroll and Pearson 1998a). In virtually all traditional statistical tests, a datum from one point in space or time is assumed to not influence or affect any another datum in the analysis obtained from a different point in space or time (independent). If, however, the data are dependent (often called autocorrelated), and many subsequent studies show that many if not most biological data are likely to be dependent, the resultant analysis may be faulty or misleading (Carroll and Pearson 2000). Many researchers now apply more appropriate statistics, such as geostatistics (Cressie 1991), in conservation biology that avoid the assumption of independence. Tiger beetles were among the first taxa using these modern analytical techniques (Carroll 1998; Pearson and Car- roll 1998; 1999; Carroll and Pearson 1998b, Pearson and Carroll 2001). In addition to pioneering statistical analyses, tiger beetles also were used in early applications of molec- ular analysis for geographical implications of conser- vation. For instance, the subdivision of lineages of the tiger beetle species, Cicindela dorsalis, in Florida between the Gulf of Mexico and the Atlantic Ocean, can be detected only with molecular markers. How- ever, the fact that species of several taxa on one side of a barrier are consistently different from those on an- other is highly significant for conservation (Pearson and Vogler 2001). These regions of distinctive genetic overlap can reflect historical events in evolutionary time (Crandall et al. 2000; Goldstein et al. 2000; Satoh et al. 2004). By incorporating an evolutionary time scale, we not only gain another valuable factor to in- clude in our conservation planning, but it also makes us aware that areas chosen for protection require man- agement goals focused not just on 10, 20 or even 100 years, but for much longer into the past as well as the future (Schwartz 1999; Barraclough and Vogler 2002). Step 5: Formation of research teams and increasing evidence of socialization, such as use of acknowledgments sections, associations of peers, and co-authored publications Conservation biology has quickly moved from single and often isolated researchers such as Leopold and Carson to a predominance of interactive researcher teams. Among tiger beetle studies, there is consider- able evidence for similar changes on a broad level, some of them apparently caused by the appearance of field guides and general books on the biology of tiger beetles beginning in the 1990s (Knisley and Schultz 1997; Leonard and Bell 1999; Acorn 200l; Choate 2003; Pearson et al. 2005; Pearson and Shetterly 2006). Before this time, only individuals with time and inter- est to search through often obscure journals and arcane terms could acquire the basic knowledge and identifi- cation skills to do research using tiger beetles. More specific evidence of socialization is in co-authored publications. In one of the first general reviews of tiger beetle biology (Pearson 1988), only 23% of the cited articles were co-authored. Twelve years later in a book on general tiger beetle biology (Pearson and Vogler 2001) 40% of its citations were co-authored. In 1969, an informal correspondence among tiger beetle enthusiasts developed into a journal called ‘‘Cicindel- a.’’ Another indicator of socialization showed advances within this highly specialized journal. In the 1970s only 2% of its articles had acknowledgments sections; in the 1980s, 26% had these sections; and in the 1990s, 83% of them did. Similarly, the complex nature of modern conserva- tion biology research necessitates more and more research teams. For instance, many modern conserva- 52 J Insect Conserv (2007) 11:47–59 123 tion biologists working on rare and endangered species now rely heavily on molecular markers (Avise 1994; Galián and Vogler 2003) to distinguish species and populations within species. The importance of con- serving intra-specific variation is reflected in the U.S. Endangered Species Act, which calls for the conser- vation of ‘‘independent population segments’’. This makes conservation of distinct populations within a species a legal requirement, and involves coordination of field biologists, laboratory technicians, lawyers, and politicians. This coordination of effort is obvious in many areas of conservation biology, and recently has also become a dominant theme in tiger beetle studies (Knisley and Hill 1992; Vogler et al. 1993; Moritz 1994; Vogler and Desalle 1994; Vogler 1998). Some promising future uses of tiger beetles have direct ramifications for conservation biology, and most of them will involve teams that are interdisciplinary. These areas include climate change (Ashworth 2001), reintroductions (Omland 2002; Brust 2002; Knisley et al. 2005), habitat reclamation (Hussein 2002), habi- tat management (Omland 2004) and location of con- servation reserves and parks (Mittermeier and Mittermeier 1997; Desender and Bosmans 1998; Andriamampianina et al. 2000; Pearson and Carroll 2001; Mittermeier et al. 2004). Step 6: Technical terminology and methodology so refined they now limit the audience that can fully comprehend it Although communication with amateurs and the public is a stated goal of the developing cadre of professional conservation biologists, growing reliance on increas- ingly complex technology and terminology, mathe- matical models, sophisticated statistics and computer programs have excluded many amateurs and even some professionals in related fields. For tiger beetles, the rapidly growing use of highly sophisticated disciplines, such as molecular biology, statistical modeling, and satellite imagery have intro- duced many technical words and concepts. This jargon, in turn, can quickly limit comprehension to a narrow array of associated professionals. As measured in terms of scientific discourse, this trend includes increasing length and number of published articles, increasing sentence complexity, use of multi-word noun phrases, as well as narrowly defined technical terms. It is also well advanced among tiger beetle workers, especially in complex fields, such as molecular studies (Galián et al. 1990; Morgan et al. 2000; Proença and Galián 2003; Goldstein and Desalle 2003; Pons et al. 2004) and mathematical modeling (Carroll and Pearson 2000; Van Dooren and Matthysen 2004). Paradoxically, although the often-growing com- plexity of terminology and methodology used in advanced studies of tiger beetles may have excluded most amateurs and many traditional taxonomists, ecologists and behavioral researchers, it appears to have attracted others. For instance, molecular biolo- gists and mathematical modelers seeking appropriate systems on which to apply their technology have used data from tiger beetles with little previous knowledge of the animals themselves. Also, when the U.S. Fish and Wildlife Service listed several tiger beetle species as endangered or threatened, economists, sociologists, foresters, politicians, land owners and members of many unrelated fields, who had little or no previous interest in these taxa, suddenly needed to know about them. At this point in the march of scientific history, the exclusion of tiger beetle amateurs from complex molecular and statistical studies, while lamentable is not debilitating. However, for conservation biology, just as the field of study reaches a high level of sci- entific rigor that knowingly will exclude many partici- pants, it simultaneously reaches a point where it must communicate with a growing number of essential participants. Many of these participants are unlikely to comprehend the message or be able to interpret the results of the increasingly complex but more reliable scientific effort. The legislators, judges, lawyers, teachers, and reporters who are critical for imple- menting policy decisions may not be able to under- stand the data and generalizations upon which they are basing their decisions. These apparently mutually exclusive goals and effort are potentially debilitating. Discussion Do the histories of tiger beetle studies and conservation biology follow the model? Both tiger beetle studies and conservation biology show patterns of change over their history consistent with the GCSPN. However, conservation biology has done so at a velocity that often blurs the progression. Studies of tiger beetles took hundreds of years to arrive at Step 6 and in the last 25 years have become greatly entwined with conservation biology. Conservation biology took less than a century to reach this level, and most of the steps were passed in the last 20 years (Primack 2002). J Insect Conserv (2007) 11:47–59 53 123 Although the GCSPN model appears to have broad relevance as shown in its application to the brief his- tory of conservation biology, the rapid advance of this field has apparently obscured some imperfections of the model along the way. Two significant questions need to be answered if the model is to be reliably ap- plied to conservation biology planning. (1) Are the steps deterministic and inevitable or are they mutable tendencies? Because the major goal of conservation biology is to protect biological diversity while providing for sustainable human needs (Primack 2002), it often seeks to change the outcome of environmental, economic and socio- logical trends, such as those associated with extinction and habitat destruction. If the general patterns of the GCSPN model represent tenden- cies that lend themselves to peremptory changes, the model can be used to anticipate problems and implement useful changes to better meet the goals of conservation biology. On the other hand, if the general patterns of the GCSPN model represent inevitable results, the changes funda- mental to conservation biology goals are unlikely to be accomplished using these general steps of science development (Myers 1989; Eldredge 1998). (2) Is each step of the model dependent on the pre- vious step, and if so, how well-developed must a step be before the subsequent step can be initi- ated and developed? For instance, academic and government support for naming and revising taxa and basic studies of natural history has been in decline for decades and is unlikely to reverse course. As crisis managers, conservation biolo- gists are often forced to make studies on taxa, natural communities and habitats that have severely incomplete foundations of knowledge, such as taxonomy and natural history (Wilson 2000; Hopkins and Freckleton 2002; Dubois 2003; Giangrande 2003). In terms of the GCSPN, the temptation is to yield to the pressures of crisis management and justify a leap from Step 1 to Step 4 or 5 with, perhaps, insufficient investment in the intermediate and supportive steps. Such a problem evidently occurred with the devel- opment of the use of bioindicators in the 1980s and 1990s. Several conservation biologists urged that these surrogate taxa be chosen carefully with predetermined ideal characteristics for a particular use and habitat or ecosystem (Brown 1991; Pearson and Cassola 1992). Unfortunately, many subsequent articles advocating taxa as bioindicators ignored or failed to adequately justify the choice of bioindicators based on predeter- mined criteria such as reliable taxonomy and basic natural history knowledge. As a result, the credibility of these poorly qualified taxa was challenged, and support of the entire concept of bioindicators quickly diminished (Lawton et al. 1998; Schwartz 1999; Andelman and Fagan 2000; Dale and Beyler 2001). In the same vein, the U.S. federal Endangered Species Act (ESA) was authorized in 1973. During its tenure, it has engendered considerable controversy, and its future is uncertain (Czech and Krausman 2001). Although property rights, conflicting economic inter- ests, and politics have contributed to many of the controversies, testimony to U.S. congressional com- mittees (Legislative Hearing on H.R. 2829 and H. R. 3705, 20 March 2002) by both conservation advocates and the Assistant Secretary for Fish and Wildlife and Parks place much of the blame for shortcomings of the ESA on poor scientific standards and lack of adequate independent scientific review of endangered species listings. For instance, in one official list of 36 species planned to be delisted in 1999 by then Secretary of the Interior, Bruce Babbitt, five species were already extinct by that time, four were based on taxonomic errors, and ten had been originally listed because of data errors. In this case 53% of these species should not have been on the endangered list in the first place, and a lack of scientific information was to blame (B. Babbitt, pers. com.). A powerful and sophisticated legislative policy assumed that conservation biology was at Step 4 or 5 in the GCSPN, even though Steps 1 and 2 were not sufficiently established to support an advance on to subsequent steps. What uses does the GCSPN provide for identifying and attaining conservation biology goals? One important role of the application of the GCSPN model to conservation biology is in providing a context so that we can focus on pertinent questions. At what points should funding agencies support specific efforts? Are there better periods than others in which to attract young recruits to maintain or increase interest in spe- cific taxa or fields such as conservation biology? Can or should dominance by a single individual or small clique be avoided? Will professional biologists exclude the expert amateurs, or will they be able to cooperate? A second use of the GSCPN is in recognizing broader philosophical problems. For instance, histori- ans of science have shown how cultural differences within national or between regional organizations 54 J Insect Conserv (2007) 11:47–59 123 often dampen paradigm changes in the general area of study (Browne 1996). Can our model illuminate factors such as this and thus avoid intellectual imperialism? Can ideas and hypotheses spread quickly throughout the network, or will resistance to change and other barriers make communication ponderous? Is there a Step 7 in our GCSPN model? Finally, these preliminary results from comparisons of tiger beetles and conservation biology highlight some specific actions that can be taken immediately. For instance in the area of communication between technical and popular audiences, a basic conservation biology goal, should or can we avoid or ameliorate Step 6? (1) Three simple changes in writing style and edito- rial format could make communication easier across a spectrum of readers. First, the abstract and summary of an article can be written in a style that simplifies complex concepts for non-profes- sionals (Gopen and Swan 1990; Knight 2003). Second, for many non-scientific readers, citations in parentheses may become a barrier that disrupts comprehension, a possibility rarely addressed or tested by scientific authors (Rudolph 2003). Using less obtrusive superscript numbers to key cita- tions is one simple change that might broaden communication. This format is already used in several prestigious journals, such as Science, Nature and Trends in Ecology and Evolution. Third, even though metaphors are central to how we think about things, especially when explaining complex concepts to the uninitiated (Short 2000), the editors of some journals, such as Conservation Biology, explicitly discourage authors from using metaphors. Encouraging the use of suitable met- aphors to enhance communication might prove more appropriate (Chew and Laubichler 2003). (2) Although administrators and professional col- leagues may demand publications in peer- reviewed journals for promotion and tenure, less prestigious methods for communicating results to the public, such as newspaper and magazine articles, books, and web sites, must receive more than a tacit blessing. (3) Even though most professional conservation biologists lack the talent or time to communicate with the public as well as Rachel Carson, Jared Diamond, Aldo Leopold, or E.O. Wilson, there are talented science writers, such as David Qua- men, Jonathan Weiner and Peter Matthiessen, who can make complex scientific writing com- prehensible and attractive to a wide range of the public who have little or no science background. Cooperating with these types of writers, even though credit may be diluted, could disseminate critical information effectively to a wider audi- ence. (4) Descriptions of new species of tiger beetles, nat- ural history observations, geographical distribu- tions, and seasonal records of occurrence and dispersion, as in many taxa, have by default been turned over largely to expert amateurs. However, not all professionals accept the resultant data as reliable. Recently the British social critics, Charles Leadbeater and Paul Miller (2004), identified a rapidly-growing involvement of ama- teurs in science from astronomy to medicine that is not fully recognized or utilized. These investi- gators are a new breed of largely self-trained experts or professional amateurs (Pro-Ams) who, using modern technology, such as the Internet, are producing significant innovations and discov- eries in a wide range of fields. Both the govern- ment and professionals need to facilitate the contributions of Pro-Ams and be prepared to share the stage with them. Conclusions As is typical of model-testing, results often reveal exceptions, unforeseen data, and other anomalies. One accepted procedure in the face of such problems, is to incorporate these unexpected results into a more gen- eralized and useful model. From results of our pre- liminary consilience tests of the GCSPN, several changes are evident that would make the model more useful. For instance, the history of tiger beetles shows that productive researchers can be working simulta- neously in several if not all the steps, especially at later times in the history of a scientific field. Thus it might be more accurate to consider the steps as benchmarks in a continuum rather than linear chronological progres- sions or irreversible advances. Also, even within well- defined taxa, amateur and professional lines of change appear to diverge into parallel lines rather than follow a single evolving line of science used in the original model (Battalio 1998) (Fig. 1). These parallel lines often have cross lines of influence and varying levels of communication. The different lines each may have their own characteristic benchmarks (Fig. 3). It is also obvious that broader fields, such as conservation biol- ogy, build on the work of contributing areas of interest and incorporate their histories rather than follow an independent disciplinary track. Thus, in these suc- J Insect Conserv (2007) 11:47–59 55 123 ceeding fields, the velocity of change along the time continuum could be expected to be faster and with entire groups of lines converging. To better understand the model, future tests are needed to clarify not only its patterns but the causes of the patterns. Consilience comparisons of the history of additional taxa or disciplines are one obvious approach. Do all taxa and fields follow the same sequence of steps? Do some histories reveal accelerated progress through certain steps and not others? Accumulated similarities and differences among these histories will provide opportunities to look for their causes. Do factors such as species numbers, their conspicuousness, economic importance, number of investigators, and level of re- search funding influence patterns and advances in the progression of steps within a field? With an under- standing of various combinations of characteristics that might cause differences in development or speed of change, we would be in a better position to understand and apply the model. Among insects, taxa such as ants, cerambycid beetles, scarab beetles, butterflies, dragon- flies, and termites would be good candidates for test organisms (New 1984; 1991; 1998; Gaston et al. 1993; Samways 1994; 2005). Comparisons of the history of fields such as wildlife biology, population genetics, and landscape ecology could also be enlightening. With some immediate solutions and the promise of even more important long range solutions made possi- ble by examinations of historical models, such as the GCSPN, we can be encouraged that conservation biology can make use of its history and learn from it. For instance, Leadbeater and Miller’s thesis indicates that with conscious effort the diverging model in Fig. 3 might morph eventually into a model with converging lines, at least between amateurs and early steps in the professional progression of changes. With improve- ments in the model and future tests of the process of science itself, we may have the best chance to develop foresight, learn from history, and better know if and what changes can be made to better reach our goals. We need not be doomed to repeat history, and even more positively, it may well be that, ‘‘We know the future only by the past we project into it’’ (Gaddis 2004). Acknowledgements We are indebted to J. Alcock, K.R. John- son, C.B. 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A model of the past using tiger beetles \(Coleoptera: Cicindelidae\) and conservation biology to anticipate the future Abstract Introduction Methods The historical model Test subjects Fig1 Results Step 1: Descriptive natural history and search for new species predominate Step 2: Now an experimental science rather than a natural history model Fig2 Step 3: Power is transferred from expert amateurs to trained professional scientists, and graduate training for employment in the field has become available Step 4: Systematics no longer dominant, and research focused more on theoretically complex issues with extensive use of graphs and statistical inference in publications Step 5: Formation of research teams and increasing evidence of socialization, such as use of acknowledgments sections, associations of peers, and co-authored publications Step 6: Technical terminology and methodology so refined they now limit the audience that can fully comprehend it Discussion Do the histories of tiger beetle studies and conservation biology follow the model? What uses does the GCSPN provide for identifying and attaining conservation biology goals? Conclusions Acknowledgements References CR1 CR2 CR3 CR4 CR5 CR6 CR7 CR8 CR9 CR10 CR11 CR12 CR13 Fig3 CR14 CR15 CR16 CR17 CR18 CR19 CR20 CR21 CR22 CR23 CR24 CR25 CR26 CR27 CR28 CR29 CR30 CR31 CR32 CR33 CR34 CR35 CR36 CR37 CR38 CR39 CR40 CR41 CR42 CR43 CR44 CR45 CR46 CR47 CR48 CR49 CR50 CR51 CR52 CR53 CR54 CR55 CR56 CR57 CR58 CR59 CR60 CR61 CR62 CR63 CR64 CR65 CR66 CR67 CR68 CR69 CR70 CR71 CR72 CR73 CR74 CR75 CR76 CR77 CR78 CR79 CR80 CR81 CR82 CR83 CR84 CR85 CR86 CR87 CR88 CR89 CR90 CR91 CR92 CR93 CR94 CR95 CR96 CR97 CR98 CR99 CR100 CR101 CR102 CR103 CR104 CR105 CR106 CR107 CR108 CR109 CR110 CR111 CR112 CR113 CR114 CR115 CR116 CR117 CR118 CR119 CR120 CR121 CR122 CR123 CR124 CR125 CR126 CR127 CR128 CR129 CR130 CR131 CR132 CR133 CR134 CR135 CR136 CR137 CR138 << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (None) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (ISO Coated) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.3 /CompressObjects /Off /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJDFFile false /CreateJobTicket false /DefaultRenderingIntent /Perceptual /DetectBlends true /ColorConversionStrategy /sRGB /DoThumbnails true /EmbedAllFonts true /EmbedJobOptions true /DSCReportingLevel 0 /SyntheticBoldness 1.00 /EmitDSCWarnings false /EndPage -1 /ImageMemory 524288 /LockDistillerParams true /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveEPSInfo true /PreserveHalftoneInfo false /PreserveOPIComments false /PreserveOverprintSettings true /StartPage 1 /SubsetFonts false /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 150 /ColorImageDepth -1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages false /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.76 /HSamples [2 1 1 2] /VSamples [2 1 1 2] >> /ColorImageDict << /QFactor 0.76 /HSamples [2 1 1 2] /VSamples [2 1 1 2] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 150 /GrayImageDepth -1 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.76 /HSamples [2 1 1 2] /VSamples [2 1 1 2] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 600 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile (None) /PDFXOutputCondition () /PDFXRegistryName (http://www.color.org?) /PDFXTrapped /False /Description << /DEU /ENU >> >> setdistillerparams << /HWResolution [2400 2400] /PageSize [2834.646 2834.646] >> setpagedevice