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1 Introduction – A Brief History of Revolutions in the Study of Insect Biodiversity 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 Insects, Arachnids, and Nematodes, Agriculture and Agri‐Food Canada, Ottawa, Ontario, Canada

John Platt (1964), in his iconic paper “Strong Hennig’s procedural framework for inferring Inference,” asked “Why should there be such relationships. The revolutions of significance rapid advances in some fields and not in in understanding biodiversity (Fig. 1.1) have, others?” The answer, he suggested, was that therefore, largely been those that enabled and “Certain systematic methods of scientific enhanced (i) the discovery process, (ii) the con­ thinking may produce much more rapid pro­ ceptual framework, and (iii) the management of gress than others.” As a corollary to Platt’s information. (1964) query, we ask “Why, within a field, should there be such rapid advances at some times and not at others?” The answer, we sug­ 1.1 Discovery gest, is “revolutions” – revolutions in thinking and technology. Perhaps the most revolutionary of all the devel­ In the study of life’s diversity, what were the opments that enabled the discovery of insect revolutions that brought us to a 21st‐century biodiversity was the light microscope, invented understanding of its largest component – the in the 16th century. The first microscopically insects? Some revolutions were taxon specific, viewed images of insects, a bee and a weevil, such as the linkage of diseases to vectors were published in 1630 (Stelluti 1630). Other (e.g., mosquitoes), which necessitated the need excellent early examples of microscope‐enabled to discover and understand species. Others illustrations of insects, such as ants, fleas, flies, included all insect taxa, such as the develop­ and even a fold‐out centerfold of a louse, were ment of light microscopy.COPYRIGHTED Some were small, featured MATERIAL in Robert Hooke’s 1665 publication, such as the invention of the Malaise trap. Some Micrographia (Neri 2011). Improvements in were mighty, such as the molecular revolution. magnification and resolution over the next two As discovery revealed an ever‐increasing wealth centuries ensured that the microscope would of biodiversity, patterns began to emerge. The continue as the primary enabler of insect biodi­ organization and explanation of these pat­ versity research. By the time light microscopy terns received quantum boosts from Carolus had achieved its theoretical limit of resolution Linnaeus’s systems of classification and nomen­ in the late 1800s, the study of insects and their clature, Charles Darwin’s natural explanation diversity had become a well‐established enter­ for species and their relationships, and Willi prise, although still largely descriptive in nature.

Insect Biodiversity: Science and Society, Volume II, First Edition. Edited by Robert G. Foottit and Peter H. Adler. © 2018 John Wiley & Sons Ltd. Published 2018 by John Wiley & Sons Ltd.

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1590 Light microscope is invented 1840s: “At the end of the abdomen are placed 1600 the anal appendages, an examination of which is

First microscopically viewed imperative for the correct discrimination of 1630 insects are illustrated species. Already, in 1842, Rambur had become 1635 France's Muséum National d'Historie Naturelle fully alive to the importance of these charac­ becomes first modern museum ters…” (McLachlan 1874, p. 6). As genitalia were 1660 analyzed for each group, the number of species increased. For example, the number of spe­ First dichotomous identification key is produced 1689 cies of black flies (Simuliidae) described from 1700 Linnaeus’s “backyard” (Fennoscandia) doubled

1720 in 1911, the year genitalia were introduced as Réné Réaumur publishes natural history taxonomic characters for the family (Lundström 1734 monographs on insect species 1911). And following the introduction of geni­ Linnaeus's Systema Naturae 10th edition talic characters for leafhoppers (Cicadellidae) in 1758 establishes beginning of modern zoological nomenclature 1922 (DeLong 1922), the discovery of new spe­ 1780 cies surged. Thomas Say publishes first With the microscope came the development comprehensive taxonomic treatment 1800 of insects in the New World of new preparation and preservation techniques (Bracegirdle 1998). Glass microscope slides, ini­ Insect genitalia are introduced as 1824 taxonomic characters tially with coverslips of mica, became dominant 1840 Chromosomes are discovered in the 1800s, and by the 20th century, coverslips Charles Darwin's On the Origin of Species of glass with standardized thickness became the 1859 is published arrangement routinely used today. The early US Taxonomic value of non-morphological federal entomologist Theodore Pergande was 1880 characters (e.g., sound production) recognized Glass microscope slides with Canada balsam using microscope slides with Canada balsam to and glass coverslips are used for insects preserve and study aphids as early as the 1870s Figure 1.1 Selected highlights in the insect biodiversity (Miller and Foottit 2017). The early choice of time line. Canada balsam, a natural product from the bal­ sam fir (Abies balsamea), as a mounting medium has ensured that slides prepared more than 100 years ago are still interpretable today. Additional developments in microscopy, includ­ When human interests collide with insects, ing those used routinely by researchers, such as science progresses. Threats to food, fiber, phase‐contrast microscopy (invented in the health, and shelter have led to dramatic leaps in early 1930s) and scanning electron microscopy discovering and understanding insect biodiver­ (first commercially available in the 1960s), sity. In the early 1800s, Rafinesque described 36 improved the ability to interpret, although species of aphids, prompted by his recognition rarely to discover, structural characters. The that these tiny insects are often deleterious light microscope, however, remains the most to their host plants (Miller and Foottit 2017). fundamental tool in insect biodiversity research. Thaddeus Harris’s splendid 1841 book and sub­ The microscope enabled an explosion of dis­ sequent expanded editions provided the vade coveries of new species and new characters that mecum for dealing with the scourges of agri­ permitted refinements in classification and culturally important insects and a foundation identification. The study of insect genitalia, for for future biodiversity exploration. In Harris’s instance, would not have been possible before (1841) words, “Some knowledge of the classifi­ the microscope. The scientific value of insect cation of insects … seems to be necessary to the genitalia was well understood by the early farmer, to allow him to distinguish his friends

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from his enemies of the insect race.” The Édouard-Gérard Balbiani discovers 1881 polytene chromosomes ­agricultural ravages of Lygus, for example, even­ 1883 Charles V. Riley suggests the existence of tually demanded deeper understanding of the what became known as cryptic species pests and helped to launch the career of noted 1893 Concept of type specimens is formalized mirid specialist Harry Knight (1917), who went 1897 Ronald Ross links malaria to mosquitoes on to describe 1345 species of plant bugs 1900 Berlese funnel is invented (Schuh 1995). 1905 Thomas H. Morgan initiates insect The year 1897 brought about a revolutionary genetic studies with Drosophila 1911 improvement to human health and ensured that Charles P. Alexander describes the first mosquitoes would become one of the taxonom­ of more than 10,000 crane fly species ically best‐known groups of insects on the 1920

planet. That was the year Ronald Ross (1897) Frits Zernike invents phase-contrast found malarial parasites in the gut of a “dappled‐ 1930 microscopy winged mosquito” (Anopheles sp.). As the focus 1934 Malaise trap is invented on vectors intensified, taxonomists bore down Ernst Mayr introduces the on the question of species and their differential 1942 biological species concept vectorial competency. Complexes of cryptic First edition of the International Code of Zoological Nomenclature is published species eventually were revealed (Coluzzi et al. 1950 2002). The genera with the most notorious vec­ Scanning electron microscopes become commercially available tors, Aedes and Anopheles, became some of the English-language version of Willi Hennig’s taxonomically best‐known mosquitoes. At least 1961 Phylogenetic Systematics is published Electronic databases, keys and computer- 75 species of Anopheles are now known to trans­ 1966 based analyses introduced mit malarial agents to humans (Foster and 1970 Electrophoresis-based taxonomy of insects Walker 2009). At a finer scale, the Anopheles becomes widely used gambiae complex includes the most efficient 1977 Fred Sanger introduces the chain-termination method for DNA sequencing 1982 malarial vectors. From genes to organisms, this Terry Erwin’s fogging of tropical canopy 1983 species complex ranks among the most taxo­ suggests 30 million insect species nomically well‐studied groups of insects. The Polymerase chain reaction (PCR) developed inevitable conclusion is that the degree of taxo­ by biochemist Kary Mullis nomic activity and sophistication is correlated Genome sequenced for Drosophila melanogaster with the severity and prevalence of disease. 2000 2003 Paul Hebert and colleagues introduce Society has always had its adventurous souls. the DNA barcode (mt COI gene) Premiere among them have been the naturalists Massively parallel sequencing who pushed into Earth’s remote frontiers, becomes readily available exploring new continents, new biomes, and 2017 new habitats to collect insects (Conniff 2011). Figure 1.1 (Continued) Tropical prospecting, in particular, yielded a torrent of new biodiversity, epitomized by Terry Erwin’s (1982) fogging of tropical canopy and cialists who devoted their lives to their central­ revolutionary suggestion that the tropical rain­ passion – a particular group of insects. Pre‐emi­ forests hold tens of millions of undiscovered nent was Charles P. Alexander (1889–1981), species. The tropics still have vast numbers of who, over a period of nearly 70 years beginning undiscovered species, but the new frontier of in 1911, described more than 10,000 species of biodiversity exploration and discovery is in the crane flies, a feat that, in his own words, “seems genome. certain it never will be done again” (Wheeler Among the chief drivers of the discovery of 1985). Examples of other productive specialists insect biodiversity have been the taxonomic spe­ include the following:

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●● Adolphe Hustache and Eduard Voss: more Origin of Species remarked: “How extremely than 9000 and more than 5000 species, stupid not to have thought of that!” (Huxley respectively, of Curculionidae (C. H. C. Lyal, 1901). personal communication); When entomologists realized that the domi­ ●● Max Bernhauer and Malcolm Cameron: 5251 nant human sense – vision – did not always and 4136 species, respectively, of Staphylinidae rule in the insect world, the opportunity was (Herman and Smetana 2001); set to discover a new realm of biodiversity: ●● Alexandre A. Girault: 4843 species of cryptic species. A pioneer in appreciating the Chalcidoidea (Noyes 2001); existence of cryptic species was C. V. Riley, who ●● Dwight DeLong: 2712 species of Cicadellidae as early as 1883 recognized that galls induced (Dmitriev 2017); on different parts of the hackberry tree repre­ ●● Hans Malicky: 2453 species of Trichoptera sented different species of psyllids (Wheeler (Morse 2017); et al. 2010). The limits of biodiversity discovery ●● José C. M. Carvalho: 2078 species of Miridae imposed by structural homogeneity were lifted (Schuh 1995). as new character sources were explored. The value of non‐visual communication signals – Trade‐offs, however, often accompanied acoustical, luminescent, and chemical – in spe­ these herculean efforts. At times, synthesis and cies discovery started to become apparent in analysis were sacrificed, and a dearth of identi­ the 1860s. But it was not until the beginning of fication keys, illustrations, or monographs hob­ the 20th century that one of the first taxonomic bled future workers faced with accessing the decisions was made on the basis of such sig­ mountain of taxonomic information needed for nals: in this case, a firefly’s flashing pattern basic identification and subsequent taxonomic (Lloyd 1990). Many new species, often isomor­ work. phic, have since been revealed on the basis of From the ubiquitous aerial net to the more songs and drumming patterns in groups as specialized variants, the development of col­ diverse as crickets, green lacewings, and stone­ lecting tools and techniques has helped to reveal flies (Alexander 1962, Stewart and Zeigler the Lilliputian world of insects. Aspirators, 1984, Henry et al. 2013). Likewise, the flashing beating sheets, kick nets, pan traps, pitfall traps, patterns of fireflies (Lloyd 1990) and chemical ultraviolet lights, and the like have become a signals such as the sex pheromones of many part of the entomologist’s standard field accou­ insects have signaled the presence of new spe­ trements. Many bear the names of their inven­ cies (König et al. 2015). tors: the Berlese funnel, invented in 1905, and The discovery of chromosomes in the early its 1918 modification, the Tullgren funnel; the 1840s (Sedgwick and Tyler 1939), and particu­ popular bulk‐collecting Malaise trap, invented larly of dipteran giant polytene chromosomes in 1934 by the eccentric René Malaise (Sjöberg by Édouard‐Gérard Balbiani in 1881 (Zhimulev 2014); and more taxon‐specific, if not obscure, et al. 2004), provided an entirely new source of tools such as the McPhail trap, dating by name characters that could reveal biodiversity hidden from 1933 but with roots in the 1890s (Steyskal beneath uniform morphology. Beginning in the 1977). Even in the current technological age, “Fly Room” of Thomas H. Morgan at Columbia with hand‐held devices enabling everything University in 1911, studies of Drosophila chro­ from geolocation of collecting sites to instanta­ mosomes have contributed epic insights into neous transmittal of information and images, the patterns and processes of insect biodi­versity. the simplest tools remain the sine qua non of the They played a central role in the evolutionary field entomologist. The simplicity of these tools synthesis (Patterson and Stone 1952), provided evokes the words of Thomas Huxley, who upon understanding of the spectacular radiation of grasping the central message in Darwin’s On the Hawaiian Drosophila (Carson and Kaneshiro

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1976), became the primary model of modern in which it is used, the ability of the user to ask genetics, and ushered in the genomics era perceptive questions and extract the insights, (Markow and O’Grady 2007). Chromosomal and the recognition of its limitations. The capa­ studies of other insects, especially black bility of bringing ever‐more powerful, less flies, chironomids, and mosquitoes, have uncov­ expensive, more accessible molecular tools to ered an abundance of biodiversity. About one‐­ bear on problems of species and their relation­ quarter of the described Nearctic species of ships is advancing at unprecedented speed. Simuliidae, for instance, were discovered through Eventually, all taxonomists and systematists studies of their polytene chromosomes (Adler might have a desktop or hand‐held laboratory et al. 2004). for the job, a trajectory reflected in the history Molecular biology – the term can be traced to of computers. 1938 (Tabery et al. 2016) – revolutionized the Other novel approaches to discovering and fields of taxonomy and systematics, and contin­ identifying species include near‐infrared spec­ ues to fuel much of biodiversity research. Early troscopy, which has its insect‐taxonomy roots electrophoretic approaches (e.g., Avise 1974) in the early 1950s (Rodríguez‐Fernández et al. reached their zenith of popularity in the 1970s 2011), and cuticular hydrocarbon analysis, and 1980s before being eclipsed by more sophis­ which began to be applied to taxonomy in the ticated techniques. The development of the 1970s (Kather and Martin 2012). Although polymerase chain reaction (PCR) technique by often profitable, these and other techniques Nobel laureate Kary Mullis in 1983, combined were not routinely adopted. General access with the introduction of chain‐termination to the necessary equipment, technology, and DNA sequencing in 1977, further enabled the expertise might limit the potential of such tech­ progress of the molecular revolution. The niques and preclude them from generating the sequencing of the entire genome of Drosophila same level of species discovery and informa­ melanogaster in 2000 (Adams et al. 2000) tional capacity enabled by simpler techniques. offered a glimpse of future possibilities. Among the most promoted, and controver­ sial, revolutions in the study of biodiversity is 1.2 Conceptual Development DNA barcoding, a natural incarnation of the universal product code that was first put into Modern biological classification and nomen­ commercial use for a pack of chewing gum in clature began officially with the tenth edition 1974 (Fox 2011). In essence, a short genetic of Systema Naturae (Linnaeus 1758). The ini­ sequence – a DNA barcode – typically the mito­ tial simplicity of the Linnaean classification chondrial cytochrome c oxidase I (COI) gene, is system for insects, based on a single source of used to discover and identify species. The COI characters – wings – was broadly appealing gene was introduced in 2003 by Paul D. N. (Sorensen 1995). This new system provided the Hebert, the “father of barcoding” (Marshall leap to a new paradigm, a starting point from 2005), and his colleagues as “the core of a global which gradual improvements could be made. bioidentification system for animals” that would The taxonomic categories established by solve the species identification problem and Linnaeus offered opportunities for investiga­ provide insights into biodiversification (Hebert tors to specialize in particular groups of et al. 2003). Major enterprises are now built insects – especially, at first, Coleoptera and around it, such as the Barcode of Life Data Lepidoptera – and generated interest in explor­ System, which has more than 5 million ing the globe in search of insects to classify sequences in its database (BOLD 2014). DNA (Sorensen 1995). barcoding has become a routine part of taxon­ Perhaps it was the diversity of insects and the omy, but like any tool, its value is in the manner variety of examples they afforded that gave

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entomologists the insight and basis to accept, 1.3 Information Management promote, and contribute to Darwin’s evolution­ ary theory, following the 1859 publication of The need to know has always demanded effi­ On the Origin of Species. Early entomologists, cient organization, management, and retrieval notably Henry Bates, Charles V. Riley, Alfred of information. These demands have produced Russel Wallace, and Benjamin Walsh, were some of the standard hallmarks of biodiversity among the most ardent supporters of evolution research, such as identification keys, mono­ by natural selection, regularly corresponding graphs, museums, and rules and guidelines. with Darwin, who considered himself an ento­ Biologists have long appreciated the utility of an mologist (Sorensen 1995). Darwin’s scientific identification key. The first dichotomous key – revolution also elevated the prestige of ento­ for plants of Britain – or at least the first proof mologists. Long considered little more than of concept, is attributed to Richard Waller in the eccentric fringe of society, entomologists 1689 (Griffing 2011). Computer‐based interac­ became some of the most qualified individuals tive multi‐access keys represent popular modern to render judgment on the big scientific issues tools that address identification needs (Dallwitz of the day (Sorensen 1995). 2000). The development of modern species con­ The lack of comprehensive treatments of taxa cepts went hand in hand with the discovery of or of regional faunas often stymied progress in new forms of insect diversity. Aristotle’s immu­ understanding insect biodiversity. Thaddeus table “eidos” (form) eventually gave way to Harris’s introductory letter to America’s father more realistic interpretations of species, most of entomology, Thomas Say, dated 7 July 1823, notably the biological species concept empha­ lamented this problem: “An ardent love of sizing reproductive isolation (Mayr 1942). Natural Science has induced me … to devote Continued debate and discussion have brought some of my leisure moments to the study of the various concepts of species into sharper Botany & Entomology; but the want of books … focus (Wheeler and Meier 2000). Yet, despite has not permitted me to make any great profi­ the insights that have come from these debates, ciency” (Weiss and Ziegler 1931). Thomas Say much of the insect‐biodiversity community subsequently provided a comprehensive treat­ continues to describe species without articulat­ ment, consolidating and making accessible the ing the concept being used. current knowledge about insect species in Willi Hennig, himself an entomologist (a dip­ North America. Earlier, Réné Réaumur in his six terist), established the methodology of phyloge­ volumes (1734–1742) of insect natural history netic systematics (cladistics), finally providing a and William Kirby and William Spence in their rigorous and testable framework for discovering four‐volume set of books (1815–1826) had done the genealogical relationships of all organisms, the same for Europe (Sorensen 1995). Say, who which Darwin’s intellectual revolution showed described more than 1000 new species, pro­ must exist. Hennig’s Grundzüge einer Theorie duced three volumes (1824–1828) on North der Phylogenetischen Systematik was written American insects (Sorensen 1995). while he was a German sanitation officer in Italy Museums, too, consolidate and organize during the Second World War and was pub­ information and make it accessible, enabling lished in 1950 (Schmitt 2013). His tour de force, the discovery of species within their cabinets. Phylogenetic Systematics – the 1966 English Museum collections are typically the sources for translation of his German book – became the revisions and monographs. The first natural his­ paradigm for biological classification. The many tory museum of the modern world might have software programs now available for phyloge­ been France’s Muséum National d’Historie netic analysis (Felsenstein 2017) are based on the Naturelle, established in 1635 (Nishida 2009). In fundamental principles presented by Hennig. America, the distinction falls to the American

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Philosophical Society in Philadelphia, with growing mass of biodiversity data, especially roots traceable to 1770; its collection later was sequence data (Bloom 2001, Attwood et al. transferred to the Academy of Natural Sciences 2011), but some aspects of information man­ of Philadelphia (Simpson 1942). Today, muse­ agement remain challenging. Premiere among ums worldwide hold roughly a billion insects these challenges is the description of new spe­ (Nishida 2009). Prospecting among these hold­ cies. The task of describing just the remaining ings is now more likely to reveal new biodiver­ species of the putatively well‐known North sity than would collecting in many parts of the American dipteran fauna will require eight full‐ world. time “Alexanders,” equivalent to 560 scientific For generations, the Linnaean nomenclatural years (Thompson 1990). Yet, given the revela­ and hierarchical classification systems have tions of DNA barcoding, which suggest an even been the heart and soul of the storage and richer North American insect fauna than previ­ retrieval system for communication about bio­ ously appreciated, the estimate of eight diversity. They have been strengthened by a set Alexanders might be too conservative (Hebert of international rules and guidelines, first for­ et al. 2016). To speed the processing of new spe­ malized and published as the International cies, proposals have been made to replace Code of Zoological Nomenclature in 1961, but descriptions with diagnoses, particularly DNA‐ their origins can be traced back to the early based diagnoses (Renner 2016), and even to 1840s. Although alternatives to the Linnaean establish new species based on photographs system have been proposed, such as the rather than physical specimens (Marshall and PhyloCode (Rieppel 2006), the Linnaean system Evenhuis 2015). Although expeditious, these remains the premiere information system for practices will need to be weighed against the the organization of life forms. sacrifice of extracting future information from The idea of type specimens, the touchstones detailed descriptions and actual physical of taxonomy, has a long history covering several specimens. centuries, although the early years were marred The universalization of the world’s scientific with loose understanding and variable use – community, beginning in the 1980s, although eventually more than 230 uses (Farber 1976). still often at the mercy of political winds, was The type concept, as understood today, was enabled by advancements of the Internet and refined with a proposal for standardization of digital age, including a personal computer on terms and definitions by Oldfield Thomas the desk of nearly every entomologist and insect (1893), including clarification of the term “type”, enthusiast. Professionals and amateurs now later given the name “holotype” by Schuchert take for granted near‐instantaneous accessibil­ (1897), and the introduction of the term ity to colleagues, digital specimens, literature, “paratype”. and online language translators. Excuses for Revolutions that led to enhanced discovery of duplication of effort, such as descriptions of the biodiversity were so successful, and the current same species (e.g., in the Nearctic and Palearctic pace of discovery has increased so dramatically regions during the Cold War), have been ren­ in recent decades, that the major impediment dered empty. we now face – the so‐called “taxonomic impedi­ ment” (Taylor 1983) – is the ability to deal with the large amount of information, particularly 1.4 Conclusions given the ever‐diminishing personnel devoted to the task. Computerized methods and tech­ The results of revolutions in the science of niques of the bioinformatics revolution that insect biodiversity are expressed in the follow­ began around the start of the new millennium ing pages by those who have benefited from, provide a mechanism to deal with the ever‐ participated in, and helped drive the study of

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insect biodiversity. We emphasize that the Avise, J. C. 1974. The systematic value of progress of biodiversity science is a cumula­ electrophoretic data. Systematic Biology 23: tive process – well stated by Courtney and 465–481. Weigmann (2016) – not a replacement process Bloom, M. 2001. Biology in silico: the of one technology for another. Future workers bioinformatics revolution. American Biology are well advised to view biodiversity holistically, Teacher 63: 400–407. from molecules to organisms, drawing on all BOLD. 2014. BOLD Systems, version 3. http:// available options to discover and interpret the www.barcodinglife.org/ [Accessed natural world. 30 December 2016]. Bracegirdle, B. 1998. Microscopical Mounts and Mounters. Quekett Microscopical Club, ­Acknowledgments London, UK. vi + 224 pp. Carson, H. L. and K. Y. Kaneshiro. 1976. Drosophila of Hawaii: systematics and We thank J. C. Morse and A. G. Wheeler, Jr, for ecological genetics. Annual Review of Ecology thought‐provoking discussions and relevant and Systematics 7: 311–345. literature; A. G. Wheeler, Jr, and Q. D. Wheeler Coluzzi, M., A. Sabatini, A. della Torre, M. A. Di for their insights on a draft of the manuscript; Deco and V. Petrarca. 2002. A polytene M. S. Caterino, C. Dietrich, and J. C. Morse chromosome analysis of the Anopheles gambiae for pointing us to the Staphylinidae, Auche­ complex. Science 298: 1415–1418. norrhyncha, and Trichoptera databases, res­ Conniff, R. 2011. The Species Seekers: Heroes, pectively; and C. H. C. Lyal for providing the Fools, and the Mad Pursuit of Life on Earth. numbers of Curculionidae described by selected W. W. Norton and Company, New York, taxonomists. New York. 464 pp. Courtney, G. W. and B. M. Wiegmann. 2016. Editorial overview: insect phylogenetics: an ­References expanding toolbox to resolve evolutionary questions. Current Opinion in Insect Science Adams, M. D., S. E. Celniker, R. A. Holt, C. A. 18: 93–95. Evans, J. D. Gocayne et al. [190 additional Dallwitz, M. J. 2000 onwards. A comparison of authors]. 2000. The genome sequence of interactive identification programs. http:// Drosophila melanogaster. Science 287: delta‐intkey.com [Accessed 28 December 2016]. 2185–2195. Darwin, C. 1859. On the Origin of Species by Adler, P. H., D. C. Currie and D. M. Wood. 2004. Means of Natural Selection, or the Preservation The Black Flies (Simuliidae) of North America. of Favoured Races in the Struggle for Life. Cornell University Press, Ithaca, New York. John Murray, London, UK. 502 pp. xv + 941 pp. + 24 color plates. DeLong, D. M. 1922. A monographic study of the Alexander, R. D. 1962. The role of behavioral North American species of the genus study in cricket classification. Systematic Deltocephalus. Ph.D. dissertation. Ohio State Zoology 11: 53–72. University, Columbus, Ohio. 129 pp. + 30 plates. Attwood, T. K., A. Gisel, N.‐E. Eriksson and E. Dmitriev, D. A. 2017. Auchenorrhyncha database. Bongcam‐Rudloff. 2011. Concepts, historical http://dmitriev.speciesfile.org/search. milestones and the central place of asp?key=Erythroneura&lng=En [Accessed bioinformatics in modern biology: a European 14 November 2017]. perspective. Pp. 3–38. In M. A. Mahdavi (ed). Erwin, T. 1982. Tropical forests: their richness in Bioinformatics: Trends and Methodologies. Coleoptera and other arthropod species. InTech, Rijeka, Croatia. Coleopterists Bulletin 36: 74–75.

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