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1

Introduction Peter H. Adler1 and Robert G. Foottit2

1 Department of Plant and Environmental Sciences, Clemson University, Clemson, South Carolina, USA 2 Canadian National Collection of , Arachnids and Nematodes, Agriculture and Agri‐Food Canada, Ottawa, Ontario, Canada

Every so often, a technical term born in the might conjure a forest, a box of ­biological community enters the popular vocab­ ­beetles, or perhaps the entire fabric of life. ulary, usually because of its timeliness, political Among scientists, the word has been defined, implications, media hype, and euphonious ability explicitly and implicitly, ad nauseum, producing to capture the essence of an issue. “Biotechnology,” a range of variants (e.g., Gaston 1996). In its “human genome,” and “stem cells” are terms as original context, the term biodiversity encom­ common in public discourse as they are in scien­ passed multiple levels of life (Wilson 1988), and tific circles. “Biodiversity” is another example. we embrace that perspective. It is “the variety of Introduced in its portmanteau form in the mid‐ all forms of life, from genes to , through 1980s by Warren G. Rosen (Wilson 1988), the to the broad scale of ecosystems” (Faith 2007). term has grown steadily in popularity. In May Biodiversity, then, is big biology, describing a 2008, the keyword biodiversity generated 17 mil­ holistic view of life. The fundamental units of lion hits on Google. Eight years later, the same biodiversity – species – serve as focal points for search produced nearly 53 million hits. studying the full panoply of life, allowing work­ Not all scientific terms are value‐neutral ers to zoom in and out along a scale from mol­ (Loike 2014). The word biodiversity, however, ecule to ecosystem. The species‐centered view has remained largely unencumbered by the eth­ also provides a vital focus for conserving life ical or political burden carried by terms such as forms and understanding the causes of declin­ “cloning” and “genetically modified organism.” ing biodiversity. Although the term biodiversity generally evokes Despite disagreements over issues ranging positive sentiment amongCOPYRIGHTED both the scientific from definitionsMATERIAL of biodiversity to phylogenetic community and the public, its meaning is often approaches, biologists can agree on four major subject to individual interpretation. Abraham points: (i) the world supports a great number of Lincoln grappled with a similar concern over insects; (ii) we do not know how many species the word “liberty.” In an 1864 speech, Lincoln of insects occupy our planet; (iii) the value of opined, “The world has never had a good defini­ insects to humanity is enormous; and (iv) too tion of the word liberty, and the American peo­ few specialists exist to inventory the world’s ple, just now, are much in want of one … but in entomofauna and to provide the expertise using the same word we do not all mean the ­necessary for conserving and sustainably using same thing” (Simpson 1998). To the layperson, its resources for societal benefit.

Insect Biodiversity: Science and Society, Volume I, Second Edition. Edited by Robert G. Foottit and Peter H. Adler. © 2017 John Wiley & Sons Ltd. Published 2017 by John Wiley & Sons Ltd.

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By virtue of the sheer numbers of individuals based partly on a view of species as structurally and species, insects, more than any other mac­ distinct from one another. Morphologically roscopic life form, command the attention of similar, if not indistinguishable, species (i.e., biologists. The number of individual insects on cryptic species) typically are not figured into Earth at any given moment has been calculated estimates of the number of insect species. If at one quintillion (1018) (Williams 1964), an putatively well‐known organisms as large as unimaginably large number on par with the crocodiles, elephants, giraffes, and whales are number of copepods in the ocean (Schubel and composites of multiple cryptic species (Wada Butman 1998) and roughly equivalent to the et al. 2003, Brown et al. 2007, Hekkala et al. number of sand grains along a few kilometers of 2011), a leap of faith is not required to realize beach (Ray 1996). The total number of insect that smaller earthlings also consist of addi­ species also bankrupts the mind. Estimates tional, hidden species. When long‐recognized offered over the past four centuries have nominal species of insects, from black to increased steadily from 10,000 species, pro­ butterflies, are probed more deeply, the repeti­ posed by John Ray in 1691 (Berenbaum, this tive result is an increase, often manyfold, in the volume), to as many as 80 million (Erwin 2004). number of species (Hebert et al. 2004, Post The number of described insect species recently et al. 2007). No zoogeographical bias in cryptic broke the 1 million mark – it currently stands at species has been detected, after correcting for 1,060,704 (Table 1.1), about 100 times the 1691 species richness and study intensity (Pfenninger estimate. Based on a figure of 1.50 million to and Schwenk 2007). We suspect that the dis­ 1.74 million described eukaryotic species in the coveries of additional cryptic species will far world (May 1998, Costello et al. 2012), insects outstrip the countering effects of synonymizing represent 61–71% of the total. existing names. The members of the class Insecta are arranged The precise number of species, however, is in 29 orders. Four of these orders – the Coleop­ not what we, as a global society, desperately tera, Diptera, , and – need. Rather, we require a comprehensive, fully account for more than 80% of all described spe­ accessible library of all volumes (i.e., species), a cies of living insects. The beetles are far in front, colossal compendium of names, descriptions, leading each of the next largest orders, the distributions, and biological information that Diptera and Lepidoptera, by a factor of more than ultimately can be transformed into a directory 2.4 (Table 1.1). A growing number of world of services. An example of societal use of plant checklists, catalogs, and inventories are available diversity provides a view of the potential treas­ online for various families and orders. Outfitted ures that insects could hold. Of the top 150 pre­ with search functions, they provide another tool scribed drugs in the United States, about 56% for handling the taxonomic juggernaut of new can be linked to discoveries in the natural plant species and nomenclatural changes. We can fore­ world (Groombridge and Jenkins 2002). The see a global registry of species in the near future great numbers of insects hold a vast wealth of that is updated with each new species or syno­ various behaviors, chemistries, forms, and nym, allowing real‐time counts for any taxon functions. Furthermore, individual insects offer (Polaszek et al. 2005). a package deal: each insect represents an eco­ The greatest concentration of insect species system of microbial life, teeming with a vast lies in tropical areas of the globe. One hectare array of species, many of which are host‐, gen­ of Amazonian forest contains more than der‐, and stage‐specific (Tang et al. 2012). Of 100,000 species of (Erwin 2004), of the 1 trillion estimated species of microorgan­ which roughly 80–85% are insects (May 1998, isms on Earth (Locey and Lennon 2016), the Stork et al. 2015). This value is more than 90% proportion specific to insects is unknown. The of the total described species of insects in the diversity, roles, and potential benefits that lie entire Nearctic region. Yet, this tropical skew is within the insect–microbiota relationship

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Table 1.1 World totals of described, living species in the 29 orders of the class Insecta, tallied May 2016.

Order* Described species References†

Microcoryphia 548 Mendes, Vol. II Zygentoma 594 Mendes, Vol. II Ephemeroptera 3,436 Morse, this volume Odonata 5,956 Morse, this volume Plecoptera 3,562 Morse, this volume Embiodea 397 Maehr and Hopkins 2016a Zoraptera 40 Maehr and Hopkins 2016b Orthoptera 26,107 Song, Vol. II Phasmatodea 2,976 Bradler, Vol. II Dermaptera 1,931 Haas, Vol. II Grylloblattodea 33 Eberhard et al., Vol. II Mantophasmatodea 19 Eberhard et al., Vol. II Blattodea 7,637 Cockroaches (5,565) + (2,072), Djernaes, Vol. II Mantodea 2,469 Otte et al. 2016 Psocoptera 5,640 Mockford, Vol. II Phthiraptera 5,239 Galloway, Vol. II Thysanoptera 6,102 ThripsWiki 2015 Hemiptera 106,971 Heteroptera (45,254; Henry, this volume) + Auchenorrhyncha (43,024; Bartlett et al., Vol. II) + Sternorrhyncha (18,693; Hardy, Vol II) Raphidioptera 248 Oswald, Vol. II Megaloptera 373 Oswald, Vol. II Neuroptera 5,813 Oswald, Vol. II Coleoptera 386,755 Bouchard et al., this volume Strepsiptera 615 Kathirithamby, Vol. II Hymenoptera 154,067 Huber, this volume Mecoptera 713 Bicha, Vol. II Siphonaptera 2,183 Galloway, Vol. II Diptera 157,971 Courtney et al., this volume Trichoptera 14,548 Morse, this volume Lepidoptera 157,761 Goldstein, this volume Total 1,060,704

* While recognizing the dynamic nature of the higher classification of the hexapods, including the combining of traditional orders (e.g., Misof et al. 2014), we follow the ordinal classification recognized by the authors of the chapters in Volumes I and II of Insect Biodiversity: Science and Society. The three orders of the Entognatha – the Collembola (ca. 8,600 species; Bellinger et al. 1996–2016), Diplura (ca. 800 species; Tree of Life Web Project 1995), and Protura (ca. 750 species; Szeptycki 2007) – are not included here with the Insecta. These three orders would add about 10,150 species, giving a total of roughly 1,071,000 species of Hexapoda in the world. † Species counts are drawn from various sources, typically from online catalogs and checklists, which are summarized by the authors of chapters in the two volumes of Insect Biodiversity: Science and Society, unless otherwise indicated.

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­represent one of the frontiers for biodiversity can imagine that in our lifetimes, automated research (Currie 2015). complete‐genome sequencing will be available To harvest the potential benefits of insects, to identify specimens as routinely as biologists taxonomists and systematists first must reveal today use identification keys. The futuristic Earth’s species and organize them appropriately handheld gadget that can read a specimen’s with collateral information that can be retrieved genome and provide immediate identification, with ease. In some respects, this is a race against with access to all that is known about the organ­ time. Biodiversity science must keep pace with ism (Janzen 2004), is no longer strictly science the changing face of the planet, particularly spe­ fiction. Yet, each new technique for revealing cies extinctions and reshufflings driven largely and organizing the elements of biodiversity by human activities such as commerce, land comes with its own set of limitations, some of conversion, and pollution. By 2007, for example, which we do not yet know. DNA‐sequence 1321 introduced species had been documented readers, for instance, will help little in identify­ on the Galapagos Islands, of which at least 37% ing fossil organisms. An integrated methodol­ are insects, including some, such as fire ogy, mustering information from molecules to (Wasmannia auropunctata and Solenopsis gem- morphology, will continue to prove its merit, inata) and the parasitic , although it is the most difficult approach for the that have had devastating consequences for the individual worker to master. Given the vast native flora and fauna (Anonymous 2007, number of insect species, however, today’s chal­ Causton and Sevilla 2008). As some species of lenges are likely to remain the same well beyond insects are being redistributed, others are disap­ the advent of handheld, reveal‐all devices: an pearing, particularly in the tropics, although the unknown number of species, too few experts, data are murky. We are forced into an intracta­ and too little appreciation of the value of insect ble bind, for we cannot know all that we are los­ biodiversity. ing if we do not know all that we have. We do Those who study insect biodiversity do so know, however, that extinction – the biodiver­ largely out of a fascination for insects; no eco­ sity crisis – is an inevitable consequence of nomic incentive is needed. But for most people, planetary abuse. Roughly 500 sq km of ’s from land developer to subsistence farmer, a per­ Amazon rainforest were deforested in 1 month sonal, typically economic, reason is required to in 2015 (Butler 2016). Using Erwin’s (2004) appreciate the value of insect biodiversity. This ­figure of 3 × 1010 individual terrestrial arthro­ value, therefore, must be translated into eco­ pods per hectare of tropical rainforest, we lost nomic gain. Papua New Guinea’s butterfly ranch­ habitat for more than one quadrillion arthro­ ing program is a spectacular example of the pods in that one point in space and time. sustainable use of insect ­biodiversity – ­conserving The urgency to inventory the world’s insect biodiversity while providing economic rewards fauna is gaining some balance through the cur­ to villagers (Insect Farming and Trading Agency rent revolution in technology, aimed in large 2008). Today’s biologists place a great deal of measure at the molecular level. Coupled with emphasis on discovering species, cataloging powerful electronic capabilities, the explosion them, and inferring their evolutionary relation­ of biodiversity information can be networked ships. Rightly so. But these activities will not, in worldwide to facilitate not only communication themselves, curry favor with the majority. We and information storage and retrieval but also believe that, now, equal emphasis must be placed itself – cybertaxonomy (sensu on developing the services of insects. We envi­ Wheeler 2007). Efforts to apply bioinformatics sion a new era, one of entrepreneurial biodiver­ on a global scale are well underway (e.g., sity that crosses disciplinary boundaries and Maddison et al. 2007, Barcode of Life Data links the expertise of insect systematists with that Systems 2014, Encyclopedia of Life 2016). We of biotechnologists, chemists, economists, engi­

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neers, marketers, pharmacologists, and others. Causton, C. E. and C. Sevilla. 2008. Latest records Only then can we expect to tap the magic well of of introduced invertebrates in Galapagos and benefits that can be derived from insects and measures to control them. Pp. 142–145. In broadly applied to society while maintaining a Galapagos Report 2006–2007, CDF, GNP and sustainable environment and conserving its bio­ INGALA, Puerto Ayora, Galapagos, . diversity. And this enterprise just might reinvig­ Costello, M. J., S. Wilson and B. Houlding. 2012. orate interest in biodiversity among aspiring Predicting total global species richness using professionals and the young. rates of species description and estimates of The chapters in this volume are written by taxonomic effort. Systematic Biology 61: biologists who share a passion for insect biodi­ 871–883. versity. The text moves from a scene‐setting Currie, C. 2015. NIH Antibiotic Discovery. overview of the value of insects through exam­ https://currielab.wisc.edu/research.php?area= ples of regional biodiversity, taxon biodiversity, NIH+Antibiotic+Discovery [Accessed 18 tools and approaches, and management and June 2016]. conservation to a historical view of the quest for Encyclopedia of Life. 2016. http://www.eol.org/ the true number of insect species. The case is [Accessed 3 May 2016]. made throughout these pages that real progress Erwin, T. L. 2004. The biodiversity question: how has been achieved in discovering and organiz­ many species of terrestrial arthropods are ing insect biodiversity and revealing the myriad there? Pp. 259–269. In M. D. Lowman and H. B. ways, positive and negative, that insects influ­ Rinker (eds). Forest Canopies. 2nd edition. ence human welfare. Although the job remains Elsevier Academic Press, Burlington, MA. unfinished, we can be assured that the number Faith, D. P. 2007. Biodiversity. In E. N. Zalta of insect‐derived benefits yet to be realized is far (principal ed.). Stanford Encyclopedia of greater than the number of species yet to be Philosophy. Metaphysics Research Lab, discovered. Stanford University, Stanford, CA. http://plato. stanford.edu/entries/biodiversity/ [Accessed 6 May 2016]. ­References Gaston, K. J. (ed). 1996. Biodiversity: a Biology of Numbers and Difference. Blackwell Science, Anonymous. 2007. CDF supports Galapagos in Oxford, UK. 396 pp. danger decision. Galapagos News Fall/Winter Groombridge, B. and M. D. Jenkins. 2002. World 2007: 2. Atlas of Biodiversity: Earth’s Living Resources Barcode of Life Data Systems. 2014. http://www. for the 21st Century. University of California boldsystems.org/ [Accessed 18 June 2016]. Press, Berkeley, CA. 256 pp. Bellinger, P. F., K. A. Christiansen and F. Janssens. Hebert, P. D. N., E. H. Penton, J. M. Burns, D. H. 1996–2016. Checklist of the Collembola of the Janzen and W. Hallwachs. 2004. Ten species in world. http://www.collembola.org [Accessed 18 one: DNA barcoding reveals cryptic species in June 2016]. the neotropical butterfly Astraptes Brown, D. M., R. A. Brenneman, K.‐P. Koepfli, J. fulgerator. Proceedings of the National Academy P. Pollinger, B. Milá, N. J. Georgiadis, E. E. of Sciences USA 101: 14812–14817. Louis, Jr., G. F. Grether, D. K. Jacobs and R. K. Hekkala, E., M. H. Shirley, G. Amato, J. D. Austin, Wayne. 2007. Extensive population genetic S. Charter, J. Thorbjarnarson, K. A. Vliet, M. L. structure in the giraffe. BMC Biology 5: 57. Houck, R. Desalle and M. J. Blum. 2011. An Butler, R. 2016. Calculating deforestation figures ancient icon reveals new mysteries: mummy for the Amazon. http://rainforests.mongabay. DNA resurrects a cryptic species within the com/amazon/deforestation_calculations.html Nile crocodile. Molecular Ecology 20: [Accessed 1 May 2016]. 4199–4215.

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