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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/318637848Biodiversity of Coleoptera: Science and SocietyChapter · July 2017DOI: 10.1002/9781118945568.ch11CITATIONS8READS6,3176 authors, including:Some of the authors of this publication are also working on these related projects:Cryptophagidae of the world View projectPaleontology of amber fossils View projectPatrice BouchardAgriculture and Agri-Food Canada233 PUBLICATIONS1,646 CITATIONSSEE PROFILEAndrew B. T. Smith116 PUBLICATIONS1,206 CITATIONSSEE PROFILEHume DouglasAgriculture and Agri-Food Canada157 PUBLICATIONS192 CITATIONSSEE PROFILEMatthew L. GimmelSanta Barbara Museum of Natural History73 PUBLICATIONS301 CITATIONSSEE PROFILEAll content following this page was uploaded by Matthew L. 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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.More species of insect have been described than of any other life form, and beetles are the great­est proportion of these. Beetles are also the most diverse group of organisms on Earth. The estimated number of described species of bee­tles is between 300,000 and 450,000 (Nielsen and Mound 1999). Our tally of 386,755 described extant species (Table 11.1) is based on figures from Ślipiński etal. (2011), whose subtotals may contain rounding errors. Many described spe­cies are known only from one locality or even one specimen (Stork 1999a, Grove and Stork 2000). Beetles are so diverse, and most species so poorly known, that even enumerating the actual number of described species remains dif­ficult, much less estimating the number of undescribed species.Erwin (1982) first proposed an estimate of the total number of beetle species on the planet, based on field data rather than on catalog num­bers. The technique used for his original esti­mate, possibly as many as 12 million species, was criticized, and revised estimates of 850,000 to 4 million species were proposed (Hammond 1995, Stork 1999b, Nielsen and Mound 1999). Some 70% to 95% of all beetle species, depend­ing on the estimate, remain undescribed (Grove and Stork 2000).The beetle fauna is not known equally well in all world regions. Yeates etal. (2003), for exam­ple, estimated that the known beetle diversity of Australia includes 23,000 species in 3265 genera and 121 families. This estimate of species is slightly lower than that reported for North America, a land mass of similar size: 25,160 spe­cies in 3526 genera and 129 families (Marske and Ivie 2003). While Marske and Ivie (2003) predicted that as many as 28,000 species could be in North America, including currently unde­scribed species, a realistic estimate of the true diversity of the little‐studied Australian beetle fauna could be,80,000 to 100,000 (Yeates etal. 2003). Both continents are probably relatively well‐studied compared with most tropical regions.Studies of beetle communities in restricted areas such as oceanic islands (Peck 2005), large administrative units (Peck and Thomas 1998, Sikes 2004, Carlton and Bayless 2007), or specific habitat types (Hammond 1990, Chandler and Peck 1992, Carlton and Robison 1998, Anderson and Ashe 2000) can provide important data on biodiversity at finer scales. Comprehensive spe­cies lists from well‐defined areas or habitat types are useful not only because they give insights about current ecosystem health and function, 11Biodiversity of ColeopteraPatrice Bouchard1, Andrew B. T. Smith2, Hume Douglas1, Matthew L. Gimmel3, AdamJ.Brunke1 and Kojun Kanda41 Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri‐Food Canada, Ottawa, Ontario, Canada2 Research Division, Canadian Museum of Nature, Ottawa, Ontario, Canada3 Invertebrate Zoology, Santa Barbara Museum of Natural History, Santa Barbara, California, USA4 Department of Biological Sciences, Northern Arizona University, Flagstaff, Arizona, USAInsect Biodiversity: Science and Society338Table 11.1 Extant families ofColeoptera, withtheestimated number ofdescribed extant world genera andspecies.Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageArchostemata — Crowsoniellidae 1 1 0 0.00Cupedidae 9 31 4 12.90Micromalthidae 1 1 1 100.00Ommatidae 2 6 1 16.67Jurodidae 1 1 0 0.00Myxophaga Lepiceroidea Lepiceridae 1 2 0 0.00Sphaeriusoidea Torridincolidae 7 60 3 5.00Hydroscaphidae 3 22 1 4.55Sphaeriusidae 1 20 10 50.00Adephaga — Gyrinidae 12 1,000 47 4.70Trachypachidae 1 6 4 66.67Rhysodidae 40 350 3 0.86Carabidae 1,500 40,000 1,660 4.15Haliplidae 4 220 39 17.73Meruidae 1 1 0 0.00Noteridae 14 250 4 1.60Amphizoidae 1 5 2 40.00Aspidytidae 1 2 1 50.00Hygrobiidae 1 5 1 20.00Dytiscidae 184 4,000 453 11.33Polyphaga Hydrophiloidea Hydrophilidae 200 3,400 302 8.88Sphaeritidae 1 5 0 0.00Synteliidae 1 7 0 0.00Histeridae 350 4,300 65 1.51Staphylinoidea Hydraenidae 42 1,600 86 5.38Ptiliidae 80 650 69 10.62Agyrtidae 8 70 4 5.71Leiodidae 342 3,700 209 5.65Silphidae 15 200 77 38.50Staphylinidae 3,500 56,000 2,078 3.71Scarabaeoidea Pleocomidae 2 50 0 0.00Geotrupidae 70 920 35 3.80Belohinidae 1 1 0 0.00Passalidae 64 800 1 0.13Trogidae 5 300 14 4.6711 Biodiversity ofColeoptera 339Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageGlaresidae 1 60 5 8.33Diphyllostomatidae 1 3 0 0.00Lucanidae 115 1,500 117 7.80Ochodaeidae 15 110 6 5.45Hybosoridae 74 570 5 0.87Glaphyridae 10 205 1 0.49Scarabaeidae 1,900 27,000 1,036 3.84Scirtoidea Decliniidae 1 2 0 0.00Eucinetidae 11 53 7 13.21Clambidae 6 170 13 7.65Scirtidae 35 800 54 6.75Dascilloidea Dascillidae 15 80 1 1.25Rhipiceridae 7 70 1 1.43Buprestoidea Buprestidae* 470 14,700 268 1.82Byrrhoidea Byrrhidae 38 430 33 7.67Elmidae 149 1,500 66 4.40Dryopidae 33 300 7 2.33Lutrochidae 1 11 0 0.00Limnichidae 37 390 3 0.77Heteroceridae 15 300 30 10.00Psephenidae 35 290 7 2.41Cneoglossidae 1 10 0 0.00Ptilodactylidae 34 500 10 2.00Podabrocephalidae 1 1 0 0.00Chelonariidae 3 250 0 0.00Eulichadidae 2 30 0 0.00Callirhipidae 9 150 0 0.00Elateroidea Rhinorhipidae 1 1 0 0.00Artematopodidae 8 45 8 17.78Brachypsectridae 1 5 0 0.00Cerophytidae 3 21 1 4.76Eucnemidae 200 1,500 29 1.93Throscidae 5 150 27 18.00Elateridae 400 10,000 1,026 10.26Plastoceridae 1 2 0 0.00(Continued)Table 11.1 (Continued)Insect Biodiversity: Science and Society340Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageDrilidae 6 120 3 2.50Omalisidae 3 8 1 12.50Lycidae 160 4,600 50 1.09Telegeusidae 3 10 0 0.00Phengodidae 31 250 12 4.80Rhagophthalmidae 6 30 0 0.00Lampyridae 110 2,200 100 4.55Omethidae 8 33 0 0.00Cantharidae 160 5,100 201 3.94Derodontoidea Derodontidae 4 30 12 40.00Nosodendridae 1 50 1 2.00Jacobsoniidae 3 20 0 0.00Bostrichoidea Dermestidae 50 1,200 44 3.67Endecatomidae 1 4 0 0.00Bostrichidae 90 570 33 5.79Ptinidae 230 2,200 107 4.86Lymexyloidea Lymexylidae 10 70 3 4.29Cleroidea Phloiophilidae 1 1 1 100.00Trogossitidae 50 600 23 3.83Chaetosomatidae 4 12 0 0.00Metaxinidae 1 1 0 0.00Thanerocleridae 7 30 1 3.33Cleridae 200 3,400 64 1.88Acanthocnemidae 1 1 0 0.00Phycosecidae 1 4 0 0.00Prionoceridae 3 160 0 0.00Mauroniscidae 5 26 0 0.00Melyridae 300 6,000 142 2.37Cucujoidea Boganiidae 5 11 0 0.00Byturidae 7 24 3 12.50Helotidae 5 107 0 0.00Protocucujidae 1 7 1 14.29Sphindidae 9 59 9 15.25Biphyllidae 7 200 2 1.00Erotylidae 260 3,500 39 1.11Table 11.1 (Continued)11 Biodiversity ofColeoptera 341Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageMonotomidae 33 250 29 11.60Hobartiidae 2 6 1 16.67Cryptophagidae 60 600 111 18.50Agapythidae 1 1 1 100.00Priasilphidae 3 11 0 0.00Phloeostichidae 5 14 1 7.14Silvanidae 58 500 15 3.00Cucujidae 4 44 8 18.18Myraboliidae 1 13 0 0.00Cavognathidae 1 9 0 0.00Lamingtoniidae 1 3 0 0.00Passandridae 9 109 0 0.00Phalacridae 51 640 31 4.84Propalticidae 2 30 0 0.00Laemophloeidae 37 430 25 5.81Tasmosalpingidae 1 2 0 0.00Cyclaxyridae 1 2 0 0.00Kateretidae 14 95 15 15.79Nitidulidae 350 4,500 211 4.69Smicripidae 1 6 0 0.00Bothrideridae 38 400 3 0.75Cerylonidae 52 450 7 1.56Alexiidae 1 50 2 4.00Discolomatidae 16 400 0 0.00Endomychidae 130 1,800 13 0.72Coccinellidae 360 6,000 306 5.10Corylophidae 30 200 23 11.50Akalyptoischiidae 1 24 0 0.00Latridiidae 28 1,000 108 10.80Tenebrionoidea Mycetophagidae 18 130 27 20.77Archeocrypticidae 10 60 0 0.00Pterogeniidae 7 26 0 0.00Ciidae 42 650 105 16.15Tetratomidae 13 150 12 8.00Table 11.1 (Continued)(Continued)Insect Biodiversity: Science and Society342Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageMelandryidae 60 420 63 15.00Mordellidae 100 1,500 130 8.67Ripiphoridae 38 400 4 1.00Zopheridae 190 1,700 74 4.35Ulodidae 14 30 1 3.33Promecheilidae 7 20 1 5.00Chalcodryidae 3 15 0 0.00Trachelostenidae 1 2 0 0.00Tenebrionidae 2,300 20,000 499 2.50Prostomidae 2 30 1 3.33Synchroidae 3 8 2 25.00Stenotrachelidae 7 19 6 31.58Oedemeridae 100 500 35 7.00Meloidae 120 3,000 80 2.67Mycteridae 29 160 2 1.25Boridae 3 4 2 50.00Trictenotomidae 2 13 0 0.00Pythidae 7 23 9 39.13Pyrochroidae 30 167 17 10.18Salpingidae 45 300 18 6.00Anthicidae 100 3,000 75 2.50Aderidae 50 900 4 0.44Scraptiidae 35 500 66 13.20Chrysomeloidea Oxypeltidae 2 3 0 0.00Vesperidae 17 75 0 0.00Disteniidae 32 336 0 0.00Cerambycidae 5,232 30,079 703 2.34Megalopodidae 30 350 6 1.71Orsodacnidae 3 40 2 5.00Chrysomelidae 2,114 32,500 1,018 3.13Curculionoidea Nemonychidae 20 70 12 17.14Anthribidae 372 3,900 50 1.28Belidae 38 375 4 1.07Caridae 4 6 1 16.67Attelabidae 150 2,500 54 2.16Table 11.1 (Continued)11 Biodiversity ofColeoptera 343Table 11.1 (Continued)Suborder Superfamily FamilyWorld generaWorld species World BINs**World percentage coverageBrentidae 400 4,000 207 5.18Curculionidae† 4,600 51,000 2,430 4.76Total 29,595 386,755 15,417 3.99Family classification is based on Bouchard etal. (2011) (except where footnoted), with generic and species numbers from Ślipiński etal. (2011). Alternative classifications to Bouchard etal. (2011) and new additions at the rank of family are highlighted in the text (in the section “Overview of Extant Taxa”). **Numbers of DNA barcoding index numbers (BINs) for each family are from the Barcode of Life Data System (BOLD: http://www.boldsystems.org) and are used to calculate the percent barcoding coverage by species. Based on data downloaded December, 2014.*Includes Schizopodidae.†Includes Dryophthoridae and Brachyceridae.but also because they can be,compared to lists generated at other periods to monitor changes over time (Howden and Howden 2001). Analysis of changes in species composition allows us to better understand human effects on ecosystems and provides evidence to guide land use and conservation decisions.To explain the great diversity of beetles is diffi­cult. One of the most important factors proposed was the development of the forewings into scle­rotized elytra (Lawrence and Britton 1994). In most beetles, the elytra cover the membranous flight wings and the abdomen. In this way, the elytra are thought to protect beetles against envi­ronmental stresses and predation (Hammond 1979). In addition, the close historical association of some of the most diverse groups of beetles, such as weevils (Curculionoidea), longhorn beetles (Cerambycidae), and leaf beetles (Chryso­melidae), with flowering plants during their own period of diversification is thought to be one of the main reasons explaining the beetles’ great diversification (Farrell 1998, Barraclough et al. 1998, McKenna et al. 2015a), but also see McKenna et al. (2015b).Patterns of beetle diversity can illustrate fac­tors that have led to the success of the group as a whole. Based on estimates for all 176 families (Table 11.1), more than 386,000 species of extant beetles have been described and are considered valid. Most species (60%) are in six megadiverse families (Fig. 11.1), each with at least 20,000 described species (in decreasing order): Staphy linidae, Curculionidae, Carabidae, Chrysome lidae, Cerambycidae, and Scarabae­idae. The smaller families of Coleoptera account for 27% of the total species in the group, and include 139 families with one to 999 described species and 28 families with 1000 to 6000 described species. So, the success of beetles as a whole is driven not only by several extremely diverse lineages, but also by a high number of moderately diverse lineages. The patterns seen today indicate that beetles went through a mas­sive adaptive radiation early in their evolution­ary history, with many of the resulting lineages flourishing through hundreds of millions of years to the present. The adaptive radiation of angiosperms may have helped to drive the diversification of beetles, as four of the six megadiverse families of beetles are primarily angiosperm feeders (Curculionidae, Chyso­melidae, Scarabaeidae, and Cerambycidae). However, even non‐phytophagous lineages of predators, scavengers, detritivores, and fungi­vores are also tremendously diverse.Beetles occur in most terrestrial and fresh­water habitats (Lawrence and Britton 1994), Insect Biodiversity: Science and Society344and a few occupy marine environments (Doyen 1976, Moore and Legner 1976). Their body length ranges from 0.3–0.4 mm (Sörensson 1997, Polilov 2008, Grebennikov 2009) to more than 17 cm. Larvae of large beetles can weigh more than 140 g, and they are the heaviest insects known (Acorn 2006). The most com­mon life‐cycle type in beetles is holometaboly, whereby individuals emerge from eggs as larvae, develop through several instars, pupate, and eventually emerge as adults. Sexual repro­duction is predominant, although partheno­genesis (i.e., production of viable, unfertilized eggs) also occurs. More specialized or unusual life cycles, which include the occurrence of active and inactive larval instars in parasi­toid species, also are known in Coleoptera (Lawrence and Britton 1994).Perhaps most people do not recognize that we live in the “age of beetles,” in which beetles are the most diverse order of organisms on Earth (The Coleopterists Society 2007). Beetles are impor­tant in most natural terrestrial and freshwater ecosystems, have a great effect on agri culture and forestry, and are useful model organisms for many types of science. A better understanding of beetle biodiversity will enhance our knowledge of the world and provide many practical applica­tions. Coverage of all aspects of beetle biodiver­sity that affect science and society is not possible in one chapter. We have therefore chosen to cover each major taxonomic group of beetles and provide some examples to illustrate important aspects of beetle biodiversity.11.1 Overview of Extant TaxaA classification of world Coleoptera was pub­lished by Bouchard etal. (2011) and is used here to present data on extant beetle diversity (Table 11.1). A brief introduction to the major beetle groups is included below, with data on sig­nificant recent knowledge advances, based on new tools and data sets. Examples of beetles from the four coleopteran suborders are included in Fig. 11.2 and overall diversity patterns in Fig.11.1. A thorough review of the diversity, distribution, biology, and relationships of each extant beetle family was published in three recent authorita­tive books (Beutel and Leschen 2005a, Leschen etal. 2010, Leschen and Beutel 2014).11.1.1 Suborders Archostemata andMyxophagaThe suborder Archostemata includes about 40 species of small to medium‐sized beetles in five families (Ommatidae, Crowsoniellidae, Figure 11.1 Examples of lesser‐known representatives of the four suborders of Coleoptera (clockwise from top left): Ptomaphagus hirtus (Tellkampf) (Polyphaga) is found only in caves in Kentucky, USA; Lepicerus inaequalis Motschulsky (Myxophaga) lives in moist sand from Mexico to Venezuela; Arthropterus wilsoni (Westwood) (Adephaga) is associated with ants under bark and logs in southeastern Australia; and Rhipsideigma raffrayi (Fairmaire) (Archostemata) is found in rotten logs in the drier forests of Madagascar. (See color plate section for the color representation of this figure.)11 Biodiversity ofColeoptera 345Micromalthidae, Cupedidae, and Jurodidae). Hörnschemeyer (2005) provides a review of the morphology and biology of the families of Archostemata, from which most of the following is taken. Most larvae develop in fungus‐infested wood, and the mouthparts of adults suggest that most species feed on plant pollen or sap, although the bionomics of members of two families (Crowsoniellidae and Jurodidae) are unknown. Ommatidae, which occur in Australia and south­ern South America, are thought to develop on subterranean dead wood. Cupedidae, with 31 species, comprise the largest and most widely distributed family, occurring on all continents except Europe and Antarctica; they generally feed on fungus‐infested wood as larvae. The fam­ily Micromalthidae includes one species that has been collected in many places throughout the world, but seems to be native to North America. The Crowsoniellidae are still known only from the original three specimens collected from soil beneath a chestnut tree in Italy, and extant Jurodidae are known only from a single specimen collected in the Far East of Russia. This is an unfortunate situation, as these two families are among the most enigmatic beetles, and fresh, DNA‐quality specimens could shed light on early beetle evolution.Myxophagans are small beetles (most less than 2.5 mm) that feed on algae or blue‐green algae in freshwater and riparian habitats (Jäch 1998, Beutel 2005). The four myxophagan families include approximately 100 species worldwide. The family Torridincolidae, the largest in the Other families24%Staphylinidae14%Curculionidae13%Carabidae10%Chrysomelidae8%Cerambycidae8%Scarabaeidae7%Tenebrionidae5%Buprestidae4%Elateridae3%Melyridae2%Coccinellidae2%Figure 11.2 Diversity of Coleoptera families (from Table 11.1). The 11 largest beetle families (each with 6000 or more described species) are shown. The remaining 154 families (each with fewer than 6000 described species) are combined into “other families,” together representing 24% of the diversity.Insect Biodiversity: Science and Society346suborder with 60 species, is exceptional in occur­ring almost exclusively,in fast‐flowing streams, spray zones of waterfalls, or hygropetric habitats (Beutel and Vanin 2005). The Hydroscaphidae occur in many aquatic habitats, including hygro­petric habitats and hot springs (Maier etal. 2010, Short etal. 2015). The Sphaeriusidae are more terrestrial than the previous families, but are usually found in close proximity to aquatic habi­tats (Beutel and Arce‐Pérez 2005). The first pos­sible larva of the poorest‐known family, Lepiceridae, was only recently described from wet leaves and flood debris along the banks of a stream in Panama (Lawrence etal. 2013); adults occur in a similar habitat.Direct effects of these two suborders on humans are minor. Examples of these include the rotten‐wood‐feeding Micromalthus debilis (“telephone‐pole beetle,” sole member of Micromalthidae), which was probably transported around the world via the timber trade (Philips 2001), and members of the family Cupedidae that live in structural timber (Neboiss 1968, 1984).11.1.2 Suborder AdephagaThe Adephaga include more than 40,000 species (Ball and Bousquet 2000, Beutel and Ribera 2005). Phylogenetic evidence supports the division of this suborder into two clades, the terrestrial Geadephaga (containing one large family, the Carabidae) and the predominantly aquatic Hydradephaga (Maddison etal. 2009, McKenna etal. 2015a). Traditionally, the Geadephaga were divided into the three families, the Carabidae, Trachypachidae, and Rhysodidae (Table 11.1), but recent evidence from molecular data suggests that the latter two families are derived members of the Carabidae. Although numerous studies have used morphological and molecular data to infer the phylogeny of the Carabidae (Maddison etal. 1999, Ribera etal. 2005), relationships among the major lineages still remain largely unresolved. Molecular data have begun to clarify relationships among the eight hydradephagan families (Ribera et al. 2002, Balke et al. 2005, McKenna et al. 2015a), and relationships within the largest family, the Dytiscidae, have been well studied (Ribera et al. 2008, Miller and Bergsten 2014). Two hydradephagan families, the Aspidytidae and Meruidae, were described from recently discov­ered beetles (Ribera et al. 2002, Spangler and Steiner 2005, respectively), highlighting the need for continued collecting (Beutel etal. 2006).The Carabidae, commonly referred to as “ground beetles,” represent the most diverse fam­ily of Adephaga, with an estimated 40,000 known species (Erwin 1991). Most carabids are carni­vores as larvae and adults, although several line­ages are herbivores (e.g., Zabrini and Harpalini), parasitoids (e.g., Brachinini and Lebiini), and spe­cialists on myxomycetes (e.g., Rhysodinae) (Arndt etal. 2005). Among carnivorous carabids, some have evolved specialized diets, such as feeding on snails (Cychrini), millipedes (Promecognathus), and ants (many Paussinae). Numerous groups have evolved to occupy unique habitats, including caves (e.g., Rhadine, Miquihuana, Dalyat, and many Trechini), endogean habitats (Anillini), and ant nests (Pseudomorphini and many Paussinae).Ground beetles are the most important adephagans in terms of anthropogenic interac­tions. A good understanding of species‐level taxonomy, especially in northern temperate regions (Lindroth 1961–1969, 1974, 1985–1986), allows scientists to use species in this diverse group as tools to address many ecologi­cal and evolutionary questions. Studies of ground beetles have enhanced our understand­ing of the ecological effects of agricultural prac­tices (Varchola and Dunn 1999, Freuler et al. 2001, Duan et al. 2006), forestry practices (Niemela etal. 1993, Magura etal. 2003), habitat fragmentation (Magagula 2003), fire manage­ment of natural habitats (Larsen and Work 2003), pollution (Freitag 1979), and other human activities. Ground beetles are also important for research on habitat conservation (Bouchard et al. 2006) and biogeography (Darlington 1943, Marshall and Liebherr 2000).Because of their predatory habits, many car­abid species are recognized as beneficial natural enemies of pestiferous insects (Lövei and Sunderland 1996, Kromp 1999). One of the 11 Biodiversity ofColeoptera 347most famous uses of carabids in biocontrol is the introduction of the European Calosoma syc-ophanta into North America to help control invasive gypsy moths. Humans also have taken advantage of the toxic nature of some carabids. The Bushmen (San people) who inhabit the Kalahari apply secretions from larvae of Lebistina spp. to their arrow tips (Koch 1958) to increase their lethality.Of the eight hydradephagan families, the Dytiscidae is by far the most species rich, with approximately 4000 described species. The rest of the families are much less diverse, with four fami­lies containing fewer than 10 species (Aspidytidae, Meruidae, Amphizoidae, and Hygrobiidae). Hydradephagans are predominantly associated with various freshwater habitats, although a few species of dytiscids are known from halobiontic environments such as saline lakes and estuaries (Balke 2005). Highly specialized groups inhabit­ing groundwater in aquifers have evolved in the Noteridae (Phreatodytes) and Dytiscidae (numer­ous genera of Hydroporinae and one species of Copelatinae). Stygobionts are known from every continent except Antarctica. The Australian fauna, which contains the highest species diver­sity, is remarkable in that it was completely unknown until the late 1990s (Watts and Humphreys 1999). Some dytiscids have lost the ability to swim and are restricted to deep, wet leaf litter in montane forests (Brancucci 1979, Brancucci and Monteith 1996). Most hydradepha­gans are predators or scavengers, although some species supplement their diet with plant material.Hydradephagans have played a smaller role in human society than have the Carabidae. Adult mosquitoes often are infected with various human disease agents, and dytiscids can help to control these disease vectors at the larval stage (Dubitskii etal. 1975, Lopez etal. 1997, Bellini et al. 2000, Lundkvist et al. 2003). An unusual traditional practice using adephagan water bee­tles recently was reported from East Africa. There, young girls collect water beetles (Gyrinidae and Dytiscidae) and hold them to their nipples to stimulate a biting reflex from these beetles. The biting and the simultaneous release of defensive secretions are locally believed to stimulate breast growth (Kutalek and Kassa 2005). Larger adult and larval dytisc­ids are also used as a source of food in numerous cultures (Ramos‐Elorduy etal. 2009).11.1.3 Suborder Polyphaga11.1.3.1 Series StaphyliniformiaThis series includes the superfamilies Hydrophiloidea sensu lato (Hydrophiloidea together with Histeroidea) and Staphylinoidea. Phylogenetic evidence is mounting that the superfamily Scarabaeoidea also should be included in this series (Hansen 1997, Beutel and Komarek 2004, Korte et al. 2004, Beutel and Leschen 2005b, Caterino et al. 2005, Hughes etal. 2006, McKenna etal. 2015a). However, it remains unclear whether the Scarabaeoidea are sister to the entire Staphyliniformia or derived within as sister to the Hydrophiloidea sensu lato (McKenna et al. 2015a, 2015b). We treat the Scarabaeiformia as a separate series to follow current classification schemes. The total num­ber of described species of Staphyliniformia is about 90,000, or roughly a quarter of all beetles and considerably more than all Vertebrata com­bined. Unlike the high diversity of the Curculionoidea, which have co‐diversified with angiosperm plants (McKenna et al. 2009), the high diversity of the Staphyliniformia seems to be a result of their repeated invasion of novel ecological niches from a generalized leaf litter‐dwelling lifestyle (McKenna etal. 2015b).The superfamily Hydrophiloidea unites the Hydrophilidae sensu,lato (Helophorinae, Epimetopinae, Georissinae, Hydrochinae, and Spercheinae are here treated as subfamilies) with the histerid group of families: Histeridae, Sphaeritidae, and Synteliidae (Archangelsky etal. 2005). The Sphaeritidae and Synteliidae include fewer than 20 species each. These families are associated with decaying organic matter, and each includes one genus distributed in North America and Eurasia. The Histeridae (350 genera and 4300 species) and Hydrophilidae (200 genera and 3400 species) are two large, cosmopolitan Insect Biodiversity: Science and Society348families that are most diverse in the tropics (Beutel and Leschen 2005a).Hansen’s (1991) work on hydrophiloid beetles, which includes taxonomic treatments down to the genus level, remains a good introduction to the family Hydrophilidae. A recent phylogeny and reclassification of the Hydrophilidae sensu stricto by Short and Fikáček (2013) features an identifica­tion key and ecological summaries of the higher taxa, and is a modern companion to the work by Hansen (1991). These beetles are primarily aquatic, occurring in a diversity of water‐based microhabitats, including those associated with waterfalls and water‐holding plants (Short and Fikáček 2013). Confirmed or suspected transi­tions to an entirely terrestrial lifestyle in leaf litter, decaying organic matter, flowers, moss, and dung are reported for most subfamilies, although the majority of terrestrial hydrophilids belong to the Sphaeridiinae (Short and Fikáček 2013). Adults are poor swimmers and normally crawl under­water, feeding mainly on plant material. Larger species prey on similar‐sized organisms. Some hydrophilids are myrmecophilous, but this is far less widespread than in other beetle families (Parker 2016). Eggs are normally stored in vari­ably shaped silk cases, which have a mast and can float; females of some Acidocerinae, Epimetopinae, and Spercheinae carry egg cases attached to their abdomens (Archangelsky etal. 2005). Larvae of most Hydrophilidae are aquatic, go through three instars, and are preda­tors; pupation, however, takes place in moist soil (Archangelsky et al. 2005). Davis (1994) considered South African dung‐inhabiting Hydrophilidae as potential biocontrol agents to be introduced against dung‐breeding flies in Australia, based on their association with herbi­vore dung.Unlike the Hydrophilidae, the Histeridae are strictly terrestrial and predatory in the adult stage, with only two larval instars (Kovarik and Caterino 2005). Fly maggots in dung, carcasses, or rotten vegetation are their most common prey. Adults of some species catch flies by hiding in crevasses in dung and seizing them with their mouthparts (Kovarik and Caterino 2005). Close associations with social insects have evolved in most of the 12 histerid subfamilies, and two of these, the Hetaeriinae and Chlamydopsinae, are predominately myrmecophilous or termitophil­ous, with unusual morphology and strong host dependency and integration (Caterino and Dégallier 2007, Parker 2016). The chapter on Histeridae by Kovarik and Caterino (2005) is a good introduction to the systematics, distribu­tion, and biology of the family. The Histeridae were used repeatedly as control agents against house flies (Musca domestica) (Kaufman et al. 2000) and other flies with dung‐inhabiting lar­vae (Davis 1994). Teretrius nigrescens, a histerid beetle, was released as a biocontrol agent against the bostrichid beetle Prostephanus truncatus to protect maize stores in equatorial Africa, although success was only moderate (Holst and Meikle 2003).The superfamily Staphylinoidea currently includes six families, and new morphological (Lawrence etal. 2011) and molecular (McKenna etal. 2015a) phylogenetic evidence suggests that the small family Jacobsoniidae (see Bostrichi­formia) also might belong here. The Hydraenidae are consistently recovered as the sister group to the Ptiliidae, as is the Agyrtidae to Leiodidae, using diverse types of phylogenetic data (Beutel and Leschen 2005b; McKenna et al. 2015a, 2015b). The Scydmaeninae and Silphidae have been shown to be derived from within the large family Staphylinidae (Ballard et al. 1998; Grebennikov and Newton 2009; Bocák et al. 2014; McKenna 2015a, 2015b), although the latter is still classified as a separate family. A consensus on the relation­ships between these higher clades has not yet been achieved (McKenna etal. 2015a).Five of these staphylinoid families are nearly worldwide in distribution, whereas the Agyrtidae are known only from northern temperate areas, except for one genus in New Zealand (Newton 1997). The family Hydraenidae comprises some 1600 described detritus‐feeding species in 42 gen­era; however, true species richness is probably higher (Jäch etal. 2005). Most hydraenid species occur in freshwater habitats; some are terrestrial and either riparian or associated with wet leaf litter. 11 Biodiversity ofColeoptera 349Their sister group, the Ptiliidae ( featherwing bee­tles), are strictly terrestrial and mycophagous and include more than 650 described species in about 80 genera (Hall 2005). The Ptiliidae include the smallest known beetles, with most species less than 1.0 mm long (Sörensson 1997). The ptiliid Scydosella musawasensis is the smallest free‐liv­ing insect, at a length of just 0.325 mm (Polilov 2015). Other remarkable features of some Ptiliidae include polymorphism in wing and eye develop­ment, association with social insects, obligate parthenogenesis, large egg size (approximately 50% of female body volume), and giant sperm (Hall 2005). The Agyrtidae represent a small fam­ily with about 70 species in eight genera that feed on decayed organic matter (Newton 1997). Their sister group, the Leiodidae, is a larger family with about 3700 species in about 342 genera (Newton 2005). Their food and habitat requirements vary, although most inhabit forests and feed on decay­ing organic matter or slime molds. Leiodids exhibit the only known case of true parasitism by beetles: Platypsyllus castori on beavers (Peck 2006). Some species of the cave‐dwelling genera Speonomus and Leptodirus have unusual larval development with only one, non‐feeding larval instar (Deleurance‐Glaçon 1963). Several genera are closely associated with colonies of ants, ter­mites, or stingless bees (Newton 2005). Newton’s (1998, 2005) works on the Leiodidae are good introductions to the diversity of this family. The influence of the Agyrtidae, Hydraenidae, Leiodidae, and Ptiliidae on humans is unclear, although they might be important decomposers of organic matter in natural and agricultural ecosystems.The Silphidae, also known as the large carrion beetles, include 200 species in 15 genera (Table11.1), most of which are efficient decom­posers of carrion, although some are preda­ceous or phytophagous (Sikes 2005). Species in the genus Nicrophorus (burying beetles) bury and actively guard small carrion to feed their larvae, with which they communicate via strid­ulation (Scott 1998). This system represents the most complex and well‐studied instance of biparental care in Staphyliniformia (in the pre­sent sense) and might be nearly 100 million years old (Cai etal. 2014).The Staphylinidae, or rove beetles, include more than 56,000 described species in 3500 gen­era across 31 subfamilies, and rival the weevils (Curculionidae) in diversity. However, within Coleoptera, rove beetles are unrivaled in mor­phological, trophic, and ecological diversity. The Staphylinidae are predominantly predaceous, with widespread instances of mycophagy and sap­rophagy for entire subfamilies (Thayer 2005). Several groups are pollen feeders, algal grazers, or parasitoids, and the genus Himalusa even feeds on the tissues of higher plants (Klimaszewski etal. 2010). Predaceous staphylinids have evolved complex rapid prey‐capture,devices, including a protrusible labium (Betz 1998) and raptorial fore­legs (Betz and Mumm 2001). Subsocial behavior in the form of larval provisioning and guarding occurs in multiple subfamilies (Ashe 1986, Wyatt 1986, Hanley and Goodrich 1995). Obligate asso­ciations with social insects have evolved more often in the Staphylinidae than in any other family of Coleoptera and the result is a remarkable array of morphological, chemical, and behavioral diver­sity, particularly in the subfamilies Aleocharinae and Pselaphinae (Parker 2016). A glowing larval staphylinid from Brazil might represent the only observation of bioluminescence in beetles outside the Elateriformia (Rosa 2010). Rove beetles are among the most commonly encountered beetles in nature, particularly in moist terrestrial habitats, and serve as indicators of human impact on natu­ral ecosystems (Work etal. 2013). As predators, some control plant pests and nuisance arthro­pods, including flies (Hu and Frank 1995) and mites (Kishimoto and Adachi 2008). As larval parasitoids and adult predators of cyclorraphous flies, members of the genus Aleochara are espe­cially well known for their biocontrol applications (Hemachandra etal. 2007). The chapter by Thayer (2005) is a thorough introduction to rove‐beetle literature.11.1.3.2 Series ScarabaeiformiaThe series Scarabaeiformia, as currently classi­fied, consists of 12 families (Pleocomidae, Insect Biodiversity: Science and Society350Geotrupidae, Belohinidae, Passalidae, Trogidae, Glaresidae, Diphyllostomatidae, Lucanidae, Ochodaeidae, Hybosoridae, Glaphyridae, and Scarabaeidae), 49 subfamilies, and 120 tribes (Smith 2006, Bouchard etal. 2011). Estimates of the number of described species in this group range from 31,000 to 35,000 (Jameson and Ratcliffe 2002, Scholtz and Grebennikov 2005). Gaston (1991) gave a conservative estimate that the actual number of species might be around 50,000 species. Scarab beetles are terrestrial, most with larvae inhabiting soil, detritus, or decaying wood. Scarab species can be found in most terrestrial habitats worldwide and are most diverse in tropical forests.Several phylogenetic analyses for taxa in the series Scarabaeiformia have been published, based on morphological characters (Howden 1982, Browne and Scholtz 1995, Browne and Scholtz 1998, Browne and Scholtz 1999, Grebennikov and Scholtz 2004), and a prelimi­nary phylogenetic analysis based on molecular characters is also available (Smith et al. 2006, Ahrens etal. 2014, McKenna etal. 2015b). The 12 families are well defined, but phylogenetic evidence for the relationships among many of these is weak or contradictory among studies. The Scarabaeidae are by far the most diverse family, containing an estimated 91% of the described species of the Scarabaeiformia (Ratcliffe etal. 2002). Two major clades occur in the family Scarabaeidae, the dung‐beetle clade and the highly diverse phytophagous scarab clade (Smith et al. 2006, Tarasov and Génier 2015). Members of the phytophagous scarab clade primarily feed on angiosperm plant foli­age, flowers, nectar, pollen, or fruit during their adult stage and roots, humus, detritus, or dead wood as larvae. The dung‐beetle clade, in their shift to dung feeding, show the second most successful trophic adaptation in the evolution of scarab beetles, after angiosperm plant feeding, when looking at total biodiversity in the group (Smith etal. 2006, Ahrens etal. 2014, Tarasov and Génier 2015). Among the other major scarab lineages are the primarily dead‐wood feeders (Lucanidae and Passalidae), carcass feeders (Trogidae), fungivores (Geotrupidae), and detritivores (Hybosoridae).Scarab beetles often have been used as a focal taxon for evolution, biodiversity, and conserva­tion research. Dung beetles, in particular, have been used in ecological and biodiversity studies (Spector 2006). Dung beetles have been used to compare habitats (Nummelin and Hanski 1989, Jankielsohn et al. 2001, Estrada and Coates‐Estrada 2002), to study diversity across land­scapes and continents (Lobo and Davis 1999, Arellano and Halffter 2003, Lobo etal. 2006), as indicator taxa of ecosystem health (Halffter and Arellano 2002, Davis etal. 2004, Quintero and Roslin 2005, Scheffler 2005), and for many other purposes.Scarab beetles are an extremely diverse, ubiqui­tous, and widely distributed group and, as such, affect humans. Hundreds of species of scarab bee­tles are nectar feeders and pollinators of diverse plant species. The benefits of dung removal and burial by dung beetles worldwide are well docu­mented from the perspective of pastureland pro­ductivity, nutrient recycling, and health (Edwards and Aschenborn 1987, Mittal 1993, Tyndale‐Biscoe 1994). In addition, dead‐wood feeding and detritivorous scarabs have a major role in nutrient and carbon cycling; burrowing scarabs play a sig­nificant part in seed burial and germination for some plant species (Vulinec 2002, Chapman and Chapman 2003), and scarabs are a major food source for many predators. Many phytophagous species are pests of turfgrass, ornamental plants, and agriculture. Numerous invasive species dam­age plants and ecosystems (Potter and Held 2002, Jackson and Klein 2006) and might outcompete native species. Some scarab species cause human health problems as carriers of disease agents (Graczyk etal. 2005).11.1.3.3 Series ElateriformiaThis group of beetles contains four superfami­lies (Dascilloidea, Buprestoidea, Byrrhoidea, and Elateroidea), 33 extant and two extinct fam­ilies (Ślipiński et al. 2011), and at least 43,000 species (Table 11.1). Of these, about 30,000 spe­cies belong to the three largest families, the 11 Biodiversity ofColeoptera 351Buprestidae, Elateridae, and Cantharidae. Elateriform beetles are found throughout the world and are most diverse in tropical regions. Chapters on Elateriformia in Beutel and Leschen (2005a), and Leschen et al. (2010) are good introductions to the classification and biology of the series and its superfamilies.A recent DNA‐based phylogenetic study (McKenna et al. 2015a) found a polyphyletic Elateriformia with weak support, including Nosodendridae but not Scirtoidea. In that study, the Dascilloidea, Byrrhoidea, and Elateroidea were monophyletic with weak support, and Buprestoidea was recovered as monophyletic with strong support. Kundrata and Bocák (2011) found that Drilidae should be reduced to a tribe of Agrypninae in Elateridae. Kundrata etal. (2014) confirmed the monophyly of Elateroidea; found new interrelationships between elateroid subfam­ilies; confirmed multiple origins of soft‐bodied­ness, neoteny, and bioluminescence; and reduced Telegeusidae to a subfamily of Omalisidae. Kundrata etal. (2016) altered the subfamily level classification of Elateridae. Douglas (2011) tested relationships in the Elateridae, using adult mor­phology, and attempted to hom*ologize morpho­logical characters for the family. The DNA‐based phylogeny of Buprestoidea by Evans etal. (2014) supported the monophyly of the Buprestoidea and of its families, with multiple origins of larval leaf‐mining behavior.Although the major known economic effects of elateriform beetles are negative (e.g., Buprestidae as forest pests and Elateridae as agricultural pests), some are beneficial. Among these, several elaterids have been introduced for biological control (of Scarabaeidae: Bianchi 1937, Clausen 1978), and Elateridae have long been known as beneficial predators of wood‐boring beetles (Craighead, 1950) and defoliators (Tostowaryk 1971). Many other predaceous lar­val and adult Elateridae, Cantharidae, and Lampyridae are expected to attack below‐ground pests and terrestrial mollusks. Similarly, the role of elateroids and buprestoids as pollina­tors is probably undervalued (but see Peter and Johnson 2005).Elateriform beetles could be,good candidates for evolutionary and ecological research. Their diverse natural history includes seven apparent independent colonizations of aquatic habitats (Lawrence 1987). They have a wider diversity of larval feeding habits than for any other series. These include feeding on dry (e.g., Buprestidae) or submerged (e.g., Elmidae) dead wood, her­bivory (including leaf mining and stem boring, e.g., Buprestidae, and moss feeding, e.g., Byrrhidae), fungivory (e.g., Elateridae, Luterek 1966), carnivory (as predators of invertebrates such as termites (Costa etal. 1992) and of verte­brates, such as sea turtle egg consumption by Elateridae (Donlan etal. 2004)), and ectoparasi­toidism (e.g., of cicadas by Rhipiceridae). Their value to evolutionary and ecological research also includes their role as primary pollinators for at least one plant species (Peter and Johnson 2005), chemical defenses, mimicry, and multiple evolu­tionary origins of bioluminescence (Branham and Wenzel 2003, Sagegami‐Oba etal. 2007).11.1.3.4 Series DerodontiformiaThe Derodontiformia include three small, quite different‐looking families of beetles, the Derodontidae, Jacobsoniidae, and Nosodendridae, with about 100 species in total. The series was only recently recognized as such, being split from the Bostrichiformia (Lawrence et al. 2010). Despite its low diversity, the group is distributed almost worldwide. The Derodontidae include about 30 species distributed in north and south temperate regions. The Jacobsoniidae, with about 20 species, occur predominantly in tropical regions, but also in New Zealand, and the Nosodendridae, with one genus (Nosodendron) and about 50 species, are distributed nearly worldwide. Habits and habitats of the group are disparate, cryptic, and often unusual; the Derodontidae are either (most genera) associated with fungi or (Laricobius) predaceous on Adelgidae (Hemiptera). The Jacobsoniidae are associated with dead wood, leaf litter, fungi, and bat guano, and the Nosodendridae are inhabit­ants of slime fluxes at tree wounds. Direct influ­ence on human beings is minimal, but species of Insect Biodiversity: Science and Society352Laricobius have shown great promise as a biologi­cal control agent of various devastating adelgid outbreaks (Lamb etal. 2006, Leschen 2011). More information on the bionomics of these families is in the respective chapters in Leschen etal. (2010).In the recent broad‐scale molecular phyloge­netic analysis of Coleoptera by McKenna etal. (2015a), each of the three families was placed in a radically different part of the tree, rendering the series grossly polyphyletic. The Derodontidae were placed with the Scirtoidea, Jacobsoniidae with Staphylinoidea, and Nosodendridae alter­natively at the base of Elateriformia or sister to Dascilloidea. Therefore, the future status of this series is in serious question.11.1.3.5 Series BostrichiformiaThe Bostrichiformia include four low‐ to medium‐diversity families. The Endecatomidae contain only four species, whereas the Bostrichidae, Dermestidae, and Ptinidae con­tain between 500 and 2200 species each. The Endecatomidae include only the genus Endecatomus, which is strictly Holarctic in dis­tribution, and feed in fruiting bodies of bracket fungi (Lawrence 2010a). The family was rela­tively recently elevated to family status out of the Bostrichidae (Lawrence and Newton 1995).The Bostrichidae are a worldwide group of about 570 species. The larvae of most members of the family are wood borers and several are major pests. The grain borers (Prostephanus and Rhyzopertha), powder‐post beetles (Lyctus and Trogoxylon), and bamboo powder‐post beetles (Dinoderus) are especially important economi­cally (Ivie 2002). One species also damages books (Hoffman 1933). The Dermestidae, with more than 1200 species, are primarily scaven­gers of dried animal and plant material, includ­ing some of the most important pests of animal‐based products, such as woolen carpets and clothes, furs, leather, and silk, whereas oth­ers are major pests of museum specimens. In addition, some carrion‐feeding dermestids are useful in forensic entomology. Some dermestids, however, have disparate life histories, Orphilus feeding in dry, fungus‐infested wood, and Thaumaglossa feeding on mantid eggs (Lawrence and Ślipiński 2010). The family Ptinidae (until recently known as Anobiidae), which is most diverse in tropical and subtropical habitats, is by far the most diverse in the series with more than 2200 species. The larvae of the Ptinidae can be separated into two groups. The first group of species bore into plant materials such as bark, seeds, dry wood, and galls, whereas the other group feeds on dried animal and plant detritus (Howe 1959, Philips 2002). Some species are associated with vertebrate nests, whereas others breed in dung. Economically important species include the death watch beetle (Anobium punc-tatum), which damages furniture, woodwork, book bindings, and other cellulose products (Peters and Fitzgerald 1996, Philips 2002). Other species of Ptinidae including the ubiquitous cig­arette beetle (Lasioderma serricorne) and drug­store beetle (Stegobium paniceum), are pests of stored products such as food commodities, wool, and products toxic to other insects, includ­ing spices, tobacco, and drugs (Howe 1959, Bousquet 1990).11.1.3.6 Series CucujiformiaThe series Cucujiformia, as defined by Lawrence and Newton (1995), includes six superfami­lies: Cleroidea, Lymexyloidea, Cucujoidea, Tenebrionoidea, Chrysomeloidea, and Curculion­oidea. An additional superfamily, Coccinelloidea, was recently defined. The superfamily Lymexyl­oidea is the smallest, containing only the family Lymexylidae. Recent morphological and molecu­lar phylogenetic studies (Lawrence et al. 2011, Bocák et al. 2014, Gunter et al. 2014, McKenna etal. 2015a) suggest that Lymexilidae have affini­ties to or should be combined with the Tenebrionoidea. Within the family, at least 60 mostly tropical species are organized in up to 12 genera (Paulus 2004). Adults of some species transmit spores of ambrosia fungi in specialized membranous cavities on their ovipositors (Wheeler 1986). Some adults (in the subfamily Atractocerinae) are longer than 60 mm and have shortened elytra resembling those of rove beetles (Staphylinidae). Some larval Lymexylidae burrow 11 Biodiversity ofColeoptera 353into wood and cultivate ambrosia fungi, which is thought to be their only larval food source. This activity causes some damage to forestry in temper­ate areas, and in the tropics Promelittomma insu-lare is a pest of coconut palms (Brown 1954). The ship‐timber worm Lymexylon navale was formerly a major pest of wooden ships (Lawrence 2010b).The superfamily Cleroidea now includes 13 families and about 10,000 species, although some authors (Majer 1994, Kolibáč 2004, Bocáková etal. 2012) advocate splitting the larg­est family, Melyridae, into several smaller fami­lies. Additionally, Robertson etal. (2015), based on phylogenetic results, transferred the families Byturidae and Biphyllidae from Cucujoidea to Cleroidea. Byturidae (24 species) occur in the Holarctic region, Southeast Asia, and southern South America; one genus, Byturus (fruitworm beetle), can be a commercial pest of Rubus (Rosaceae) fruits. The Biphyllidae (200 species) occur throughout the world except in New Zealand; species seem to feed on cryptic fungi under bark of dead trees and in leaf litter. Most larvae of the Cleroidea seem to be predators or scavengers, whereas the adults are primarily predators or pollen feeders. The larvae of some species pursue and attack wood‐boring insects inside their tunnels, benefitting forestry. The Acanthocnemidae have one extant species, Acanthocnemus nigricans, which is native to Australia and is attracted to fires and recently burnt wood (Lawrence and Leschen,2010). The Chaetosomatidae include three genera (Kolibáč 2004) and about 12 species in New Zealand and Madagascar; they are predatory (Lawrence etal. 1999). The Cleridae are perhaps the most con­spicuous elements of the superfamily, with about 200 genera and 3400 described species, many of them brightly colored and diurnal. Most are predatory as adults and larvae, including some (e.g., Thanasimus spp.) that are important in controlling outbreaks of forest pests such as bark beetles (Curculionidae: Scolytinae) (Kolibáč 2010). Mauroniscidae are a small, rarely encoun­tered family occurring in the New World; some feed in flowers, probably on pollen, but almost nothing further is known about the bionomics of the group, which was recognized only recently as a family apart from the Melyridae (Majer 1994). The Melyridae (sensu lato, including Rhadalidae, Dasytidae, and Malachiidae sensu Majer 1994, Majer 2002, and Bocáková et al. 2012) have about 6000 species (Bocáková etal. 2012). They are an extremely diverse group of mostly soft‐bodied beetles that frequent flower­ing plants as adults, feeding primarily on pollen. The Metaxinidae recently were established as a separate family (Kolibáč 2004) to accommodate one predatory, sooty‐mold‐inhabiting species from New Zealand, Metaxina ornata. There is just a single species in the family Phloiophilidae, Phloiophilus edwardsii, which is found in Europe; the larvae feed on fungi (Lawrence etal. 1999). The Phycosecidae include only the beach‐dwelling genus Phycosecis, with four species endemic to Australia, New Caledonia, New Hebrides, and New Zealand (Leschen and Beutel 2010). The Prionoceridae have at least three gen­era in the Oriental and Afrotropical regions, and are pollen feeders as adults (Lawrence et al. 1999). The Thanerocleridae, with 30 species in seven mostly tropical genera, are known from all continents except Antarctica (Kolibáč 1992, Kolibáč 2004). The heterogeneous family Trogossitidae, which contains fewer than a thou­sand species, is the third largest in the Cleroidea, and has a similar nearly worldwide distribution (Kolibáč 2005).The superfamily Cucujoidea has received a major reorganization in recent years (Robertson etal. 2015). Until recently, it included about 20,000 species in about 1500 genera in 35 families (Pakaluk etal. 1994), but now it includes only 25 families with about 915 genera and 10,950 species. Major recent changes include removal of the Byturidae and Biphyllidae to the Cleroidea (Robertson et al. 2015), carving of the Coccinelloidea out of the Cucujoidea (Robertson etal. 2015), splitting of the Cybocephalidae from the Nitidulidae (Cline etal. 2014), and combina­tion of the Propalticidae with the Laemophloeidae (McElrath et al. 2015). The newly defined Cucujoidea consist of the following families (McElrath et al. 2015, Robertson et al. 2015): Insect Biodiversity: Science and Society354Agapythidae, Boganiidae, Cavognathidae, Cryptophagidae, Cucujidae, Cybocephalidae, Cyclaxyridae, Erotylidae, Helotidae, Hobartiidae, Kateretidae, Laemophloeidae, Lamingtoniidae, Monotomidae, Myraboliidae, Nitidulidae, Passandridae, Phalacridae, Phloeostichidae, Priasilphidae, Protocucujidae, Silvanidae, Smicripidae, Sphindidae, and Tasmosalpingidae. The most diverse family in the group is the Nitidulidae, which boast an extraordinary range of habits and habitats, including fungus feeders, leaf miners, sap feeders, carrion feeders, pollen feed­ers, fruit feeders, seed predators, inquilines in ant and termite nests, and even an aquatic mosquito‐feeding species (Jelínek etal. 2010). One species, the small hive beetle (Aethina tumida), is a pest of honeybee colonies and recently spread to the United States (1998), Canada (2002), and Australia (2002) from southern Africa (Hood 2004). Closely related to the Nitidulidae are the Kateretidae and Smicripidae, which feed in flowers; the latter fam­ily is restricted to the New World. The small family Cybocephalidae are important predators of ster­norrhynchous Hemiptera, especially scale insects, and have been part of biocontrol programs. The Monotomidae are often found under bark or in decaying plant material; some are mycophagous, whereas others (e.g., Rhizophagus) are effec­tive predators of destructive bark beetles (Curculionidae: Scolytinae). Most Laemo­phloeidae occur under bark, with a few possibly being predaceous on bark beetles, and some (especially Cryptolestes species) being important stored‐product pests. The Passandridae are ectoparasitic as larvae on wood‐inhabiting beetles and Hymenoptera. The Silvanidae contain many economically important pests of stored products, including species of Oryzaephilus, Nausibius, Cathartus, and Ahasverus, but are otherwise mostly found under bark of dead trees. Most cucu­joid species are of little direct relevance to human economy because they are associated with decay­ing vegetation and fungi. The Boganiidae are pol­len feeders associated with cycads and Myrtaceae, and occur only in Australia, New Caledonia, and South Africa (Escalona et al. 2015). The Cavognathidae are unique in exclusively inhabiting bird nests; they occur in Australia, New Zealand, and South America. The Agapythidae and Cyclaxyridae are both sooty‐mold‐inhabiting bee­tles endemic to New Zealand. Most Cryptophagidae are associated with microfungi, and occur in a wide variety of habitats worldwide. Larvae of the small but widely distributed family Cucujidae can be predaceous. The Erotylidae are a large group of often colorful forms; most feed on either fungi (especially macrofungi) or plants, although a few are cycad feeders; some are even pests of stored grain. The Hobartiidae are fungus feeders that occur only in Australia and Chile. The Lamingtoniidae, Myraboliidae, and Tasmosalpingidae are all Australian endemic fam­ilies, occurring on fungi or under bark. The Phalacridae occur worldwide; some are strongly associated with smut or rust fungi, whereas others (especially Olibrus) develop in flower heads. The Phloeostichidae are known from Australia, south­ern South America, and the Palearctic region. The Priasilphidae occur in south temperate regions and feed on microfungi in forested areas. The Protocucujidae occur in Australia and southern South America, and their bionomics are obscure. The Sphindidae are distributed worldwide and feed only on slime molds. The various cucujoid family chapters in Leschen et al. (2010), from which most of the previous information was taken, serve as entry points into detailed morphological and taxonomic treatments.The superfamily Coccinelloidea, newly erected by Robertson etal. (2015), contains the group of families previously referred to as the “Cerylonid Series” of Cucujoidea. Before Robertson etal. (2015), it contained nine fami­lies, but these were fractured into 15 families in that publication: Akalyptoischiidae, Alexiidae, Anamorphidae, Bothrideridae, Cerylonidae, Coccinellidae, Corylophidae, Discolomatidae, Endomychidae, Eupsilobiidae, Euxestidae, Latridiidae, Murmidiidae, Mycetaeidae, and Teredidae. Most of the families are relatively small, but one family, the Coccinellidae, domi­nates in terms of diversity, with about 6000 spe­cies in 350 genera worldwide (Ślipiński and Tomaszewska 2010). Coccinellidae have a 11 Biodiversity ofColeoptera 355marked impact on human activities, with most species being predators, which can be useful for control of a variety of plant pests, mostly ster­norrhynchous Hemiptera, with a few feeding on spider mites. Harmonia axyridis, the widely introduced multicolored Asian lady beetle, eco­logically displaces native coccinellids, and is a nuisance pest entering human houses in search of hibernation places, and contaminating wine grapes (Pervez and Omkar 2006). Still other species (in the subfamily Epilachninae) are plant,feeders as larvae and adults, and eat mostly squashes, beans, and nightshades, on which they can be serious pests. Most of the other families are primarily obscure, cryptic fungus or slime‐mold feeders, including the recently elevated (from Latridiidae) family Akalyptoischi idae (Lord et al. 2010), which occurs only in western North America; the family Alexiidae, which occurs only in the west­ern Palearctic; the families Anamorphidae, Cerylonidae, Corylophidae, Euxestidae, and Latridiidae, eachof which occur worldwide; the family Discolomatidae, which occurs pantropi­cally; the small family Mycetaeidae, which has been introduced around the world and can be a pest of moldy stored products (Tomaszewska 2010); the family Murmidiidae, with a similar cosmopolitan distribution and habitat to the Mycetaeidae (Ślipiński and Lawrence 2010); the Bothrideridae, which include specialized para­sitoids of coleopteran and hymenopteran wood‐boring insects (Ślipiński etal. 2010); the Eupsilobiidae, which are poorly known, but can occur in association with ants (Tomaszewska 2010); and the Teredidae, a widely distributed group generally of fungus feeders and com­mensals in bark beetle tunnels (Ślipiński etal. 2010).The superfamily Tenebrionoidea includes 28 families (Table 11.1), 19 of which are minor, each with only 5 to 500 described species (Archeocrypticidae, Boridae, Melandryidae, Mycetophagidae, Mycteridae, Promecheilidae, Prostomidae, Pterogeniidae, Pyrochroidae, Pythidae, Ripiphoridae, Salpingidae, Scraptiidae, Stenotrachelidae, Synchroidae, Tetratomidae, Trachelostenidae, Trictenotomidae, and Ulodidae). The small family Trachelostenidae has recently been downgraded to a tribe within Tenebrionidae by Matthews and Lawrence (2015). The relationships and diversification patterns of beetles in this morphologically challenging super­family have benefited from analyses of molecular data sets (Gunter etal. 2014; Kergoat etal. 2014a, 2014b). Typical feeding habitats include under dead tree bark or in fungi, decaying wood, and lit­ter. Larvae of the Ripiphoridae are unusual beetle endoparasites of immature insects (bees, wasps, and co*ckroaches). The Mycetophagidae include minor stored‐product pests (perhaps feeding on fungus‐infested commodities) and a known vec­tor of disease agents (Bousquet 1990, Hald etal. 1998). One Hemipeplus (Mycteridae) species is a pest of palms (Lepesme 1947, Zelazny and Pacumbaba 1982).The families Aderidae, Anthicidae, Ciidae, Meloidae, Mordellidae, Oedemeridae, and Zopheridae are moderately diverse within the Tenebrionoidea, each with 550 to 3000 described species. A large proportion of these beetles develop in dead wood, fungi, and leaf lit­ter and are uncommonly encountered by people (Leschen et al. 2010). Nacerdes melanura (Oedemeridae) is a pest of wooden marine structures and archeological timber (Pitman et al. 1993, 2003). Beetles of the families Meloidae and Oedemeridae produce antipreda­tor secretions called cantharidins that can harm humans or kill domesticated animals. A few species in the family Anthicidae prey on the eggs and small larvae of pests and are useful bio­logical control agents (McCutcheon 2002).By far the most diverse tenebrionoid family is the Tenebrionidae, with nearly 20,000 described species. This family is expected to include many undescribed species, especially in tropical regions. Recent studies of tropical groups have greatly increased the number of known species (Bouchard 2002). Different groups of Tenebrionidae have evolved remarkable adaptations to survive in some of the most extreme habitats in the world. The production of glycerol‐like compounds in adults of Upis ceramboides enables their survival under Insect Biodiversity: Science and Society356tree bark during prolonged freezing periods to −50 °C (Miller 1978, Lundheim and Zachariassen 1993). Conversely, unusual morphological and behavioral adaptations in other tenebrionids ena­ble them to survive in the hottest deserts (Cloudsley‐Thompson 1964, McClain etal. 1985, Zachariassen 1991). Detailed studies of these adaptations might yield important applications for humans. Tenebrionids are a major faunal compo­nent of most deserts and have a significant role in food webs (Crawford et al. 1993). Shifts in the abundance and range of xerophiles, versus moist‐forest species, have been used as indicators of cli­mate change (Geisthardt 2003). Several pest tenebrionid species have been transported across the world for centuries with stored products (Andres 1931, Chaddick and Leek 1972). These beetles cause billions of dollars in direct and indi­rect losses to stored products every year. One major pest, Tribolium castaneum, is among the most‐studied insects. Similar to adephagans, many tenebrionids produce antipredator secre­tions containing a combination of repulsive agents (e.g., quinones and phenols), wetting agents, and spreading agents (e.g., hydrocarbons) or other compounds with unknown purposes (Hurst etal. 1964, Tschinkel 1975). These repulsive secretions also have antibacterial or antifungal properties (Blum 1981). Some endangered Tenebrionidae species, especially flightless species, require habi­tat conservation to avoid extinction.The superfamily Chrysomeloidea is a group of seven beetle families (Beutel and Leschen 2005a) with more than 50,000 species arranged in the cerambyciform (Cerambycidae, Disteniidae, Oxypeltidae, and Vesperidae) and chrysomeli­form (Chrysomelidae, Megalopodidae, and Orsodacnidae) lineages. Because they are almost exclusively phytophagous, many are economically important as plant pests or biological agents against weeds. The families Cerambycidae (long­horn beetles) and Chrysomelidae (leaf beetles) are by far the largest in this superfamily and include more than 30,000 and 32,000 species, respectively (Table 11.1). Reid (2014) recently reviewed phylogenetic analyses of the chrysomel­iform lineage, and Reid (2000) provided a key to subfamilies based on adults and larvae. Švácha et al. (1997) provided the most recent phyloge­netic hypothesis of the cerambyciform lineage based on larvae. Although most species in the Cerambycidae are winged and have arboreal hab­its, specialized adaptations to ground‐dwelling habits are known in the flightless genus Dorcadion and related genera (Plavilstshikov 1958). The remaining five families combined include fewer than 1000 species. The Disteniidae have more than 300 species occurring in all major biotic regions, except Australia, Europe, and New Zealand (Lawrence et al. 1999, Švácha and Lawrence 2014b). The Oxypeltidae include three large and brightly colored South American spe­cies in two genera (Lawrence et al. 1999). The Vesperidae have nearly 80 species in the warmer regions, among which the soil‐dwelling larvae of Philus antennatus are notorious pests of pine plantations in China (Švácha and Lawrence 2014a). The pantropical Megalopodidae include some 350 species, and the Orsodacnidae have approximately 40 species in three genera in the Nearctic, Neotropical, and Palearctic regions (Clark and Riley 2002, Lawrence and Ślipiński 2014).The classification of the herbivorous super­family Curculionoidea has been, and continues to be, in a state of flux, based on recent works (Bouchard etal. 2011, Löbl and Smetana 2011, Löbl and Smetana 2013, Leschen and Beutel 2014). This superfamily currently contains six smaller families with fewer than 4500 described species each (Attelabidae, Anthribidae, Belidae, Brentidae, Caridae, and Nemonychidae), as well as the second most diverse family of beetles, the Curculionidae (Table 11.1). Most species in this superfamily have the mouth on a distinctive cylindrical extension of the head. This rostrum is used to prepare cavities in plant tissues for egg deposition. The evolution of the rostrum is thought to be a,reason for the weevils’ success because it allows them to exploit plant tissues that other beetles cannot access (Anderson 1995).Members of the six smaller families of Curculionoidea generally are associated with plants that predate flowering plants (e.g., conifers, 11 Biodiversity ofColeoptera 357cycads, and ferns) (Farrell 1998). These weevils are also important pollinators for their host plants (Kevan and Baker 1983). The anthribid Araecerus fasciculatus (coffee‐bean weevil) feeds on dried plant materials and has become a seri­ous pest of stored products such as coffee and cocoa beans (Childers and Woodruff 1980, Bousquet 1990). Some species of the subfamily Apioninae (Brentidae) are specific to one or sev­eral host plants and have been introduced in new regions as biological tools to control invasive weeds (O’Brien 1995).The family Curculionidae is one of the most diverse groups of organisms, with more than 50,000 species described and thousands more to be formally described. Their close association with flowering plants is a main factor explaining their great diversity. Curculionids feed on plants of any terrestrial or freshwater habitat, and on any plant tissue, from roots to seeds (Anderson 2002). Because of these characteristics, curcu­lionids are important economically. Some spe­cies are pests of agriculture and forestry, including about half of the species of important plant pests; others feed on stored‐plant com­modities. Curculionids have been used to con­trol populations of invasive plants (O’Brien 1995). The subfamilies Scolytinae and Platypodinae (considered separate families until recently) are specialized in exploiting woody plants (Anderson 2002). Most either feed on the phloem of the inner bark of their hosts or on symbiotic ambrosia fungi that they cultivate in the wood. About 100 species in these two sub­families are major forestry pests.11.2 Overview of Fossil TaxaThe first beetle‐like fossils originated in the Early Permian (approximately 280 mya); how­ever, the true Coleoptera appeared in the Triassic (approximately 230 mya) (Grimaldi and Engel 2005, Lawrence and Ślipiński 2013). A large amount of data on fossil Coleoptera has been published and significant new discoveries (i.e., description of new taxa up to the rank of family) are published on a regular basis (Kukalová‐Peck and Beutel 2012; Bai etal. 2012, 2013; Kirejtshuk and Nel 2013; Kirejtshuk and Azar 2013; Kirejtshuk etal. 2016). The four sub­orders of extant Coleoptera (Archostemata, Myxophaga, Adephaga, and Polyphaga) and 63% of extant beetle families are known from the fossil record (Smith and Marcot 2015). Ślipiński et al. (2011) reported a total of 230 genera and 600 fossil species of beetles, and Smith and Marcot (2015) established a database of 5553 beetle fossil occurrences from 221 local­ities. A list of beetle families only represented by fossils is given in Table 11.2.11.3 Societal Benefits and Risks11.3.1 Beetles ofEconomic Importance11.3.1.1 Negative Effects ofBeetlesAgriculture Beetles have a major effect on the world’s agriculture. Hundreds of species of bee­tles, including many in the families Chrysomelidae, Curculionidae, Elateridae, and Scarabaeidae, feed on crops and ornamental plants as adults and larvae (Campbell et al. 1989). At least one species probably attacks each cultivated seed‐plant species around the world. One of the best‐studied pests is the boll weevil (Anthonomus grandis grandis), which harms cotton. This species is native to tropical and subtropical America, but has been estab­lished in the United States since the late 1800s (Burke etal. 1986). In the past 100 years, yield losses and control costs against this species have been estimated at more than US$22 billion (Kaplan 2003). A boll weevil eradication pro­gram that started in 1983 (Dickerson et al. 2001), coordinated by the US Department of Agriculture and state agencies, has resulted in the eradication of the weevil from more than 80% of the cotton‐production area in the United States.Blister beetles can contaminate animal feed. These meloid beetles are sometimes abundant in hay fields and possess antipredator cantharidin Insect Biodiversity: Science and Society358Table 11.2 Coleoptera families represented by extinct taxa only, withnumber ofdescribed genera andspecies (fromŚlipiński etal. 2011).Suborder Superfamily Family World genera World speciesProtocoleoptera Tshekardocoleoidea Tshekardocoleidae 13 18Labradorocoleidae 1 1Oborocoleidae 2 2Permocupedoidea Permocupedidae 11 30Taldycupedidae 6 18Ademosynidae 12 29Permosynidae 3 14Archostemata — Triadocupedidae 8 16Magnocoleidae 1 1Obrieniidae 5 7Myxophaga Asiocoleoidea Asiocoleidae 1 1Tricoleidae 3 6Rhombocoleoidea Rhombocoleidae 5 10Schizophoroidea Schizophoridae 20 36Catiniidae 5 6Schizocoleidae 4 30Adephaga — Tritarsusidae 1 1Triaplidae 1 2Colymbotethidae 2 2Parahygrobiidae 1 1Coptoclavidae 14 32Liadytidae 2 6Polyphaga Scarabaeoidea Coprinisphaeridae 4 9Pallichnidae 2 3Scirtoidea Elodophthalmidae 1 2Mesocinetidae 4 7Elateroidea Praelateriidae 2 2Berendtimiridae 1 1Lasiosynidae 4 7Cleroidea Parandrexidae 3 5Cucujoidea Sinisilvanidae 1 1Curculionoidea Ulyanidae 2 3Total 145 309Taxa are ordered following the classification of Bouchard etal. (2011). New family‐group additions published since 2011 are highlighted in the text (in “Overview of Fossil Taxa”).11 Biodiversity ofColeoptera 359compounds. Cattle, emu, goats, horses, and sheep have become ill or died after consuming alfalfa contaminated by dead blister beetles (Capinera etal. 1985). The incidence of this con­dition, termed cantharidiasis, has increased recently due to the common practice of hay con­ditioning, which crushes the beetles inside the hay and incorporates them into the bales (Capinera etal. 1985, Blodgett etal. 1992).Beetles are important vectors of pathogens to crops and livestock (Harris 1981). One species, the striped cucumber beetle (Acalymma vit-tatum), is a specialist herbivore of the cucumber family Cucurbitaceae. The association of these beetles with the wilt‐inducing plant bacterium Erwinia tracheiphila often leads to major losses (Garcia‐Salazar et al. 2000). Management of bacterial wilt relies heavily on vector manage­ment (Ellers‐Kirk and Fleisher 2006) through field and laboratory studies of the bionomics of the striped cucumber beetle and the effects of varying agricultural practices. The lesser meal­worm beetle (Alphitobius diaperinus) com­monly colonizes poultry houses, feeding on shed feathers, droppings, and spilled poultry meal, as well as on dead, dying, or newly born chicks (Harris 1966, Kumar 1988, Watson etal. 2001). Although the beetles have a minor bene­ficial effect in consuming chicken mites and house fly maggots, the benefits are more than offset by their role in the transmission of dis­ease‐causing bacteria and fungi such as Campylobacter, Metarhizium, and Salmonella (Davies and Wray 1995, Alves et al. 2004, Strother etal. 2005).Hinton (1945) stated that the Coleoptera was the most important order of insects attacking stored products. More than 600 species of bee­tles are associated with stored products around the world. Although many of these eat the stored products, others feed on associated fungi or animals (Bousquet 1990). Some of the most problematic groups of beetles in stored prod­ucts are the Dermestidae, Chrysomelidae, Curculionidae, and Tenebrionidae. The red flour beetle (Tribolium castaneum), a frequently encountered pest of stored products, can feed on many products in the larval and adult stages. Because of its economic importance and ease of rearing, the red flour beetle is used extensively as a tool in genetics research.Museum Collections Museums assemble millions of specimens for research and public,displays. Beetles, especially the family Dermestidae, threaten the world’s museum collections. Species of Anthrenus, Dermestes, and Thylodrias com­monly attack and destroy preserved animals such as pinned insects and preserved pelts and skins, as well as cultural artifacts that incorporate feathers, fur, hair, leather, wool, and other animal deriva­tives (Armes 1988, Campbell etal. 1989). The con­trol of museum beetle pests is becoming increasingly difficult because pesticides and fumi­gants used in the past can cause health problems for museum workers. Museum‐pest research is now focusing largely on preventive conservation (Goldberg 1996).Forestry Many species of Coleoptera are highly relevant to humans through their activity as forest pests. The assemblage of forest pests varies among countries. Up to 45% of the annual wood volume grown in Sweden was estimated to be lost to two bark beetles: Tomicus piniperda and Ips typographus (Eidmann 1992). Dendroctonus fron-talis, feeding on loblolly pine (Pinus taeda), killed about 40,000 trees with a value exceeding $10 million in the southeastern United States in 18 months (Strom etal. 2004). These and many other species together cause tree mortality levels simi­lar to those caused by fire or commercial harvest.Weevils of the subfamilies Scolytinae and Platypodinae (Curculionidae), with 5812 and 1463 species, respectively (Wood and Bright 1992), are the most infamous forest pests. Phloem‐feeding, tree‐killing bark beetles con­stitute most of the diversity of the Scolytinae. Female bark beetles normally burrow in phloem and lay eggs, from which larvae hatch and con­struct further feeding burrows. Phloem of dying or freshly killed trees is the typical environment Insect Biodiversity: Science and Society360for bark beetles. Many bark beetles use chemi­cal signals (pheromones) to schedule a syner­getic attack by many individuals on one tree, thus overcoming its resistance, killing it, and making it suitable for beetle reproduction (Paine et al. 1997). Other species overcome tree defenses with the aid of pathogenic fungi (Farrell etal. 2001).One notorious phloem feeder is, however, not a scolytine weevil, but rather the emerald ash borer, Agrilus planipennis, a jewel beetle (Buprestidae). This species, native temperate east Asia, recently was accidentally introduced to and became estab­lished in North America. The larvae feed on phloem of ash trees (Fraxinus spp.) and cause widespread decline and mortality of the trees. The species was first detected in July 2002 in Detroit (Michigan, USA) and Windsor (Ontario, Canada). The cumulative death toll of ash trees is estimated to be as high as 15 million, many of which are high‐value urban trees (Poland and McCullough 2006). The genus Agrilus is among the most species‐rich genera of all organisms, with 2767 species (Bellamy 2003), and many pose a risk of introduction and loss of forest ecosys­tems around the world.Phloem‐feeding scolytine beetles are among the species most easily introduced accidentally to new regions, primarily through dunnage and wood packaging used in international trade. Island countries such as New Zealand are par­ticularly vulnerable to introductions because of their isolated and mostly endemic native biota. Some 103 species of scolytine weevils were inter­cepted at New Zealand’s borders between 1950 and 2000 (Brockerhoff et al. 2006), including high‐risk species such as Dendroctonus pondero-sae and I. typographus. The mainland Eurasian species Dendroctonus micans was first recorded in Wales in 1982 and then spread throughout the United Kingdom (Gilbert etal. 2003). The North American species Dendroctonus valens was introduced to China in 2001 and currently infests more than 0.5 million hectares of pine forest (Cognato etal. 2005). The pine shoot bee­tle (T. piniperda) is a Palearctic bark beetle that was recently introduced to North America. Since its discovery in Ohio in 1992, this beetle has become established in Ontario and Quebec and in 13 states in the northcentral and north­eastern United States (Morgan et al. 2004). Haack (2006) reviewed the history of bark‐ and wood‐boring beetles introduced to the United States over the past 20 years and concluded that two exotic Buprestidae, five Cerambycidae, and 18 Scolytinae species have been introduced and are expanding their ranges in North America.Although most forestry‐important Coleoptera species are phloem feeders, many beetles also tunnel through sapwood, weakening or killing the host tree. Ambrosia weevils of the Scolytinae and Platypodinae have evolved a type of agricul­ture, cultivating ambrosia fungi in their burrows (Farrell etal. 2001). The Asian longhorn beetle Anoplophora glabripennis emerged recently as a top‐profile beetle in the news in North America and Western Europe. In the early 1990s, this species hitchhiked with international trade and infested urban forests in the United States and Canada, rapidly killing eudicot trees of many species. This prompted government agencies to cut down thousands of trees in Massachusetts, New Jersey, New York, Ohio, and Toronto in an ongoing effort to eradicate the species. The genus Anoplophora has some 35 other species in Northwest Asia, which were revised by Lingafelter and Hoebeke (2002). Their study indicates that other Anoplophora species also could become introduced pests overseas.Forestry in the southern hemisphere is based mainly on the hardwood Eucalyptus trees, which also have an associated complex of beetle pests. The Christmas beetle Anoplognathus chloro-pyrus (Scarabaeinae) and the leaf beetles Paropsis atomaria, Paropsis charybdis, and some other taxonomically poorly known paropsines (Chrysomelinae) are significant defoliators of commercially grown eucalypts in Queensland, Australia (Johns etal. 2004, Nahrung 2004).Many highly destructive fungal forest patho­gens are transmitted by beetles. Black‐stain root disease of conifers in western North America, for example, is caused by the ophiostomatoid fungus Leptographium wageneri. Spores of this 11 Biodiversity ofColeoptera 361species were detected in 37% of bark beetles in California, with individual beetles carrying zero to more than 100,000 spores (Schweigkofler etal. 2005). The mountain pine beetle (D. pon-derosae) and its mutualistically associated blue‐stain fungi (Grossmannia spp.) have benefitted from unusually high winter temperatures, which have lowered larval mortality in recent years. These factors created the largest beetle epi­demic in Canada’s history, primarily killing lodgepole pines (Pinus contorta var. latifolia) in British Columbia (Kim et al. 2005). This phe­nomenon of beetle–fungus mutualistic associa­tion is not restricted to bark and ambrosia beetles, but has evolved independently in a few other beetle taxa.An understanding of the biodiversity of tree‐associated beetles is relevant to the world forest industry. Only a few groups, such as the European and North American bark and ambrosia beetles, are adequately known taxonomically. Elsewhere, most tree‐living beetle species remain inade­quately known.A list of Coleoptera species of high economic importance was compiled from various sources and is presented in Table 11.3.11.3.1.2 Positive Effects ofBeetlesBiological Control of Weeds Plant‐feeding bee­tles, primarily of the families Curculionidae and Chrysomelidae, have been used success­fully to control the spread of invasive alien plant species throughout the world. Weed seeds have been transported accidentally throughout the world for centuries, and the trend is increasing with global trade (Mason et al. 2002). Once established in a new area, weeds often spread quickly, primarily because of the absence of insect herbivores associated with them. Biological control of weeds, or the introduction and manipulation,of natural ene­mies to reduce the spread of invasive weeds, is often used as a pesticide‐free control method. Careful studies of the biology of beetles in the weed’s country of origin and destination coun­try are necessary for safe and effective control (O’Brien 1995, Rea 1997, Lindgren etal. 2002). Recent success stories include control of houndstongue (Cynoglossum officinale) using a weevil (De Clerck‐Floate etal. 2005) and con­trol of purple loosestrife (Lythrum salicaria) using two species of leaf beetles (Lindgren etal. 2002).Dung Removal One cow can produce more than 9000 kg of solid waste every year (Fincher 1981). The accumulation of dung from agriculture pol­lutes waterways, reduces pasture quality, and creates microhabitats for the development of flies and other pests. Dung beetles (Scarabaeidae and Geotrupidae) promote manure decomposi­tion by burying it in the ground for larval food. They annually avert economic losses to the United States of $0.38 billion by accelerating burial of livestock feces (Losey and Vaughan 2006). Dung beetles have been imported into countries, such as Australia, where native spe­cies did not remove an increase of livestock dung (Hughes et al. 1975). They also provide a vital ecosystem service in urban environments, where they use dog dung, thereby reducing stream pol­lution and improving nutrient‐poor soils (Wallace 2005).Pollination Beetles, along with other insect groups, are crucially important in the pollina­tion of cultivated and wild plants. Pollination by beetles is often referred to as cantharophily. The Coleoptera are considered to be the most primi­tive pollinators (Kevan and Baker 1983). Beetles have been associated with flowering plants over millions of years, leading in some cases to the evolution of specialized structures in the host‐plant flowers and in their beetle pollinators (Barth 1985, Fenster et al. 2004). Thirty‐four families of flowering plants contain at least one primarily beetle‐pollinated species (Bernhardt 2000). In some cloud forests, more than 45% of palms and herbs are beetle‐pollinated (Seres and Ramirez 1995). A decline in native and non‐native pollinators worldwide was attributed to pesticide use, habitat loss, plant biodiversity loss, and poor agricultural practices (Allen‐Wardell etal. 1998). If this trend continues, we Table 11.3 Pest Coleoptera species ofhigh economic concern, based oncurrent CABI, USDA‐APHIS, Japan‐MAFF, NAPIS, CFIA, andEPPO (A1 andA2) lists.SuperfamilyFamilySubfamilyGenusSpeciesAuthor and year[Adephaga]CarabidaeCarabinaeZabrustenebrioides(Goeze, 1777)ScarabaeoideaScarabaeidaeCetoniinaeCheirolasiaburkei(Westwood, 1843)Chilolobaacuta(Wiedemann, 1823)Pachnodamarginata peregrinaKolbe, 1906interrupta(Olivier, 1789)Protaetiaauripes(Hope, 1831)bipunctata(Kraatz, 1889)celebica(Wallace, 1867)cretica(Kraatz, 1880)cuprea(Fabricius, 1775)himalayanaMikšić, 1987milaniAntoine and Pavicevic, 1994noxJanson, 1881speciosa(Adams, 1817)speciosissimus(Scopoli, 1786)Smaragdesthesafricana(Drury, 1773)DynastinaeDiloboderusabderus(Sturm, 1826)GolofaeacusBurmeister, 1847Heteronychusarator(Fabricius, 1775)OryctesagamemnonBurmeister, 1847boas(Fabricius, 1775)monoceros(Olivier, 1789)rhinoceros(Linnaeus, 1758)PapuanauninodisPrell, 1912woodlarkiana(Montrouzier, 1855)Podischnusagenor(Olivier, 1789)Xylotrupesgideon(Linnaeus, 1767)Table 11.3a (Continued)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearMelolonthinaeAmphimallonmajale(Razoumowsky, 1789)solstitiale(Linnaeus, 1758)Costelytrazealandica(White, 1844)HolotrichiadisparilisArrow, 1916mindanaonaBrenske, 1893serrata(Fabricius, 1787)Leucopholisirrorata(Chevrolat, 1841)lepidophoraBlanchard, 1851Maladeracastanea(Arrow, 1913)MelolonthahippocastaniFabricius, 1801melolontha(Linnaeus, 1758)Phyllophagasmithi(Arrow, 1912)spp.—RutelinaeAdoretussinicusBurmeister, 1855spp.—versutusHarold, 1869AnomalasulcatulaBurmeister, 1844Exomalaorientalis(Waterhouse, 1875)PopilliajaponicaNewman, 1838spp.—BuprestoideaBuprestidaeAgrilinaeAgrilusanxiusGory, 1841biguttatus(Fabricius, 1777)hyperici(Creutzer, 1799)auroguttatusSchaeffer 1905planipennisFairmaire, 1888sulcicollisLacordaire, 1835BuprestinaeChrysobothrisfemorata(Olivier, 1790)Melanophilafulvoguttata(Harris, 1829)ChrysochroinaeSphenopteragossypiiCotes, 1891(Continued)Table 11.3 (Continued)BostrichoideaDermestidaeMegatominaeTrogodermagranariumEverts, 1898BostrichidaeBostrichinaeAmphiceruscornutus(Pallas, 1772)Heterobostrychusaequalis(Waterhouse, 1844)Sinoxylonunidentatum(Fabricius, 1801)DinoderinaeProstephanustruncatus(Horn, 1878)LymexyloideaLymexylidaeMelittommatinaeMelittommainsulareFairmaire, 1893CucujoideaLaemophloeidae—Cryptolestescapensis(Waltl, 1834)NitidulidaeCarpophilinaeCarpophilusobsoletusErichson, 1843CillaeinaeMeligethesaeneus(Fabricius, 1775)NitidulinaeAethinatumidaMurray, 1867Stelidotageminata(Say, 1825)CoccinellidaeCoccinellinaeChilocoruskuwanaeSilvestris, 1909Chnootribasimilis(Thunberg, 1871)ColeomegillamaculataTimberlake, 1943Epilachnaborealis(Fabricius, 1775)chrysomelina(Fabricius, 1775)varivestisMulsant, 1850Harmoniaaxyridis(Pallas, 1773)Henosepilachnaelaterii(Rossi, 1794)vigintioctopunctata(Fabricius, 1775)ElateroideaElateridaeAgrypninaeConoderusrufangulus(Eschscholtz, 1829)DendrometrinaeLimoniuscalifornicus(Mannerheim, 1843)ElaterinaeAgrioteslineatus(Linnaeus, 1767)sputator(Linnaeus, 1758)ustulatus(Schaller, 1783)Melanotuscommunis(Gyllenhal, 1817)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearSasajiscymnustsugae(Sas,aji and McClure, 1997)Stethoruspunctum(LeConte, 1852)ChrysomeloideaCerambycidaeCerambycinaeAeolesthessarta(Solsky, 1871)Aromiabungii(Faldermann, 1835)Callidiellumrufipenne(Motschulsky, 1862)villosulum(Fairmaire, 1900)CerambyxcerdoLinnaeus, 1758ChloridolumalcmeneThomson, 1865thomsoni(Pascoe, 1859)Chlorophorusannularis(Fabricius, 1787)strobilicolaChampion, 1919Cordylomeratorrida(Fabricius, 1775)Enaphalodesrufulus(Haldeman, 1847)Euryphaguslundi(Fabricius, 1793)Hesperophanescampestris(Faldermann, 1835)Hylotrupesbajulus(Linnaeus, 1758)Oemonahirta(Fabricius, 1775)PhoracantharecurvaNewman, 1840semipunctata(Fabricius, 1775)Placosternusdifficilis(Chevrolat, 1862)Plagionotusarcuatus(Linnaeus, 1758)Stromatiumbarbatum(Fabricius, 1775)Trichoferuscampestris(Faldermann, 1835)Xylotrechusaltaicus(Gebler, 1836)chinensis(Chevrolat, 1852)colonus(Fabricius, 1775)namanganensisHeyden, 1885pyrrhoderusBates, 1873quadripesChrevrolat, 1863(Continued)rufiliusBates, 1884spp.—Xystroceraglobosa(Olivier, 1795)LamiinaeAnoplophorachinensis(Forster, 1771)glabripennis(Motschulsky, 1854)spp.—AnthoresleuconotusPascoe, 1869ApomecynabinubilaPascoe, 1858Aprionacinerea(Chevrolat, 1852)germari(Hope, 1831)japonica(Thomson, 1878)Batocerarufomaculata(De Geer, 1775)Bixadussierricola(White, 1858)DectestexanusLeConte, 1862Desmiphorahirticollis(Olivier, 1795)Lagocheirusaraneiformis(Linnaeus, 1767)MonochamusalternatusHope, 1842carolinensis(Olivier, 1792)galloprovincialis(Olivier, 1795)saltuarius(Gebler, 1830)sartor(Fabricius, 1787)scutellatus(Say, 1824)spp.—sutor(Linnaeus, 1758)urussovi(Fischer von Waldheim, 1805)Oberealinearis(Linnaeus, 1761)oculata(Linnaeus, 1758)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)Psacotheahilaris(Pascoe, 1857)SaperdacalcarataSay, 1824candidaFabricius, 1787carcharias(Linnaeus)perforata(Pallas, 1773)Steirastomabreve(Sulzer, 1776)LepturinaeEncyclopscaerulea(Say, 1826)PrioninaePrionuscalifornicusMotschulsky, 1845laticollis(Drury, 1773)SpondylidinaeArhopalusferus(Mulsant, 1839)Tetropiumcastaneum(Linnaeus, 1758)fuscum(Fabricius, 1787)gracilicorneReitter, 1889ChrysomelidaeBruchinaeAcanthoscelidesobtectus(Say, 1831)BruchuslentisFrölich, 1799Callosobruchusanalis(Fabricius, 1781)rhodesianus(Pic, 1902)Caryedonserratus(Olivier, 1790)CassidinaeBrontispalongissima(Gestro, 1885)CoelaenomenoderaelaeidisMaulik, 1920Dicladispaarmigera(Olivier, 1808)gestroi(Chapuis, 1877)Odontotadorsalis(Thunberg, 1805)PromecothecacumingiiBaly, 1858Trichispasericea(Guérin‐Méneville, 1844)ChrysomelinaeGonioctenafornicataBrüggemann, 1873Leptinotarsadecemlineata(Say, 1824)Mesoplatyscincta(Olivier, 1790)MicrothecaochrolomaStål 1860(Continued)PhaedonbrassicaeBaly, 1874CriocerinaeCriocerisasparagi(Linnaeus, 1758)duodecimpunctata(Linnaeus, 1758)Liliocerislilii(Scopoli, 1763)Oulemamelanopus(Linnaeus, 1758)EumolpinaeTypophorusnigritus(Fabricius, 1801)GalerucinaeAcalymmavittatum(Fabricius, 1775)Alocyphabimaculata(Jacoby, 1903)Aulacophorafoveicollis(Lucas, 1849)indica(Gmelin, 1790)Cerotomatrifurcata(Forster, 1771)Chaetocnemabasalis(Baly, 1877)confinisCrotch, 1873pulicariiaMelsheimer, 1847DiabroticabalteataLeConte, 1865barberiSmith and Lawrence, 1967speciosa(Germar, 1824)undecimpunctataMannerheim, 1843virgifera virgiferaLeConte, 1868Epitrixucumeris(Harris, 1851)asciatalatchley, 1918imilarisentner, 1944ubcrinita(LeConte, 1857)uberisentner, 1944Exosomalusitanicum(Linnaeus, 1767)Luperomorphaxanthodera(Fairmaire, 1888)MadurasiaobscurellaJacoby, 1896Medythiaquaterna(Fairmaire, 1880)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)OothecabennigseniWeise, 1900mutabilis(Sahlberg, 1829)PhyllotretachotanicaDuvivier, 1892cruciferae(Goeze, 1777)striolata(Illiger, 1803)Pyrrhaltaviburni(Paykull, 1799)CurculionoideaAttelabidaeRhynchitinaeCaenorhinusmarginellus(Faust, 1898)BrentidaeApioninaeIschnopterapionvirens(Herbst, 1797)BrentinaeArrhenodesminutus(Drury, 1773)Cylasbrunneus(Olivier, 1790)formicarius(Fabricius, 1798)puncticollisBoheman, 1833BrachyceridaeBrachycerinaeBrachycerusspp.—ErirhininaeLissorhoptrusoryzophilusKuschel, 1952DryophthoridaeOrthognathinaeRhinostomusbarbirostris(Fabricius, 1775)RhynchophorinaeCosmopolitessordidus(Germar, 1824)Diocalandrafrumenti(Fabricius, 1801)taitense(Guérin‐Méneville, 1834)Metamasiushemipterus(Linnaeus, 1758)spp.—Odoiporuslongicollis(Olivier, 1807)Rhabdoscelusobscurus(Boisduval, 1835)Rhynchophorusferrugineus(Olivier, 1791)palmarum(Linnaeus, 1758)ScyphophorusacupunctatusGyllenhal, 1838Sphenophorusvenatus(Say, 1832)Temnoshoitanigroplagiata(Q,uedenfeldt, 1888)Tryphetusincarnatus(Gyllenhal, 1838)CurculionidaeConoderinaeCopturusaguacataeKissinger, 1957CossoninaeCaulophilusoryzae(Gyllenhal, 1838)(Continued)CryptorhynchinaeCryptorhynchusmangiferae(Fabricius, 1775)Elytroteinussubtruncatus(Fairmaire, 1881)Euscepespostfasciatus(Fairmaire, 1849)MitrastethusbaridioidesRedtenbacher, 1868Sternochetusmangiferae(Fabricius, 1775)frigidus(Fabricius, 1787)CurculioninaeAnthonomusbisigniferSchenkling and Marshall, 1934eugeniiCano, 1894grandisBoheman, 1843pomorum(Linnaeus, 1758)quadrigibbusSay, 1831signatusSay, 1832Curculiocaryae(Horn, 1873)elephas(Gyllenhal, 1836)nucumLinnaeus, 1758Orchestesalni(Linnaeus, 1758)CyclominaeGonipterusgibberusBoisduval, 1835scutellatusGyllenhal, 1833ListroderescostirostrisSchönherr, 1826subcinctusBoheman, 1842Listronotusbonariensis(Kuschel, 1955)oregonensis(LeConte, 1857)EntiminaeBlosyrusasellus(Olivier, 1807)Diaprepesabbreviatus(Linnaeus, 1758)famelicus(Olivier, 1790)splengleri(Linnaeus, 1767)Geniocremnuschilensis(Boheman, 1842)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)Hypomecessquamosus(Fabricius, 1792)Naupactusleucoloma(Boheman, 1840)xanthographus(Germar, 1824)Otiorhynchusarmadillo(Rossi, 1792)corruptor(Host, 1789)cribricollisGyllenhal, 1834dieckmanniMangano, 1979ligustici(Linnaeus, 1758)meridionalisGyllenhal, 1834pseudonotusApfelbeck, 1897singularis(Linnaeus, 1767)sulcatus(Fabricius, 1755)ovatus(Linnaeus, 1758)rugosostriatus(Goeze, 1777)Pantomoruscervinus(Boheman, 1840)PhlyctinuscallosusSchoenherr, 1826Premnotrypeslatithorax(Pierce, 1914)suturicallusKuschel, 1956vorax(Hustache, 1933)Pseudocneorhinusbifasciatus(Roelofs, 1880)Sitonacylindricollis(Fåhraeus, 1840)discoideusGyllenhal, 1834hispidulus(Fabricius, 1777)humeralisStephens, 1831TanymecusdilaticollisGyllenhal, 1834HyperinaeHyperapostica(Gyllenhal, 1813)LixinaeBothynoderespunctiventris(Germar, 1824)Hypolixustruncatulus(Fabricius, 1798)LixusjunciiBoheman, 1835(Continued)MesoptiliinaePissodescastaneus(De Geer, 1775)harcyniae(Herbst, 1795)nemorensisGermar, 1824strobi(Peck, 1817)terminalisHopping, 1920MolytinaeAlcidodesdentipes(Olivier, 1790)ConotrachelusaguacataeBarber, 1924nenuphar(Herbst, 1797)perseaeBarber, 1919spp.—HeilipuslauriBoheman, 1845Hylobiusabietis(Linnaeus, 1758)transversovittatus(Goeze, 1777)PlatypodinaeCrossotarsusexternedentatus(Fairmaire, 1850)squamulatusChapuis, 1865DiapusminutissimusSchedl, 1969pusillimusChapuis, 1865quinquespinatusChapuis, 1865Dinoplatypusagnatus(Schedl, 1969)biuncus(Blandford, 1896)cavus(Strohmeyer, 1913)chevrolati(Chapuis, 1865)cupulatulus(Schedl, 1941)cupulatus(Chapuis, 1865)forficula(Chapuis, 1865)luniger(Motschulsky, 1863)pallidus(Chapuis, 1865)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)pseudocupulatus(Schedl, 1935)uncatus(Browne, 1980)Euplatypuscompositus(Say, 1824)hintzi(Schaufuss, 1897)parallelus(Fabricius, 1801)Genyocerusborneensis(Browne, 1980)diaphanus(Schedl, 1939)pendleburyi(Schedl, 1936)spinatus(Browne, 1980)Megaplatypusmutatus(Chapuis, 1865)Oxoplatypusquadridentatus(Olivier, 1795)Platypusapicalis(White, 1846)curtusChapuis, 1865cylindrus(Fabricius 1792)excedensChapuis, 1865geminatusChapuis, 1865jansoniChapuis, 1865koryoensis(Murayama, 1930)parallellus(Fabricius, 1801)porcellusSchedl, 1942pseudocurtusSchedl, 1935quercivorus(Murayama, 1925)subdepressus(Schedl, 1935)westwoodiChapuis, 1865Treptoplatypussolidus(Walker, 1859)ScolytinaeAncipitispunctatissimus(Eichhoff, 1880)AnisandrusmaicheStark, 1936(Continued)dispar(Fabricius, 1792)Arixyleboruscanaliculatus(Eggers, 1923)granifer(Eichhoff, 1878)granulifer(Eggers, 1923)hirsutulusSchedl, 1969imitator(Eggers, 1927)mediosectus(Eggers, 1923)rugosipesHopkins, 1903Cnestusmutilatus(Blandford, 1894)Coccotrypessubcribrosus(Blandford, 1896)CryphaluslatusEggers, 1929CryptoxyleborussubnaevusSchedl, 1937Crypturguscinereus(Herbst, 1794)Cyclorhipidionsexspinatum(Schedl, 1935)DendroctonusadjunctusBlandford, 1897armandi(Tsai and Li, 1959)brevicomisLeConte, 1876frontalisZimmerman, 1868micans(Kugelann, 1794)ponderosaeHopkins, 1902pseudotsugaeHopkins, 1905rufipennis(Kirby, 1837)valensLeConte, 1857Dolurguspumilus(Mannerheim, 1843)DryocoetesconfususSwaine, 1912affaber(Mannerheim, 1852)Eccoptopterusspinosus(Olivier, 1795)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)EuwallaceafornicatusEichhoff,1868destruens(Blandford, 1896)Gnathotrichussulcatus(LeConte, 1868)retusus(LeConte, 1868)Hylastesater(Paykull, 1800)Hylesinusvarius(Fabricius, 1775)aculea,tusSay, 1826Hylurgopinusrufipes(Eichhoff, 1868)Hylurgopspalliatus(Gyllenhal, 1813)rugipennis(Mannerheim, 1843)Hylurgusligniperda(Fabricius, 1787)Hypothenemushampei(Ferrari, 1867)Ipsamitinus(Eichhoff, 1872)calligraphus(Germar, 1824)cembrae(Heer, 1836)confusus(LeConte, 1876)duplicatus(Sahlberg, 1836)grandicollis(Eichhoff, 1868)hauseri(Reitter, 1894)leconteiSwaine, 1924paraconfususLanier, 1970pini(Say, 1826)plastographus(LeConte, 1868)sexdentatusBörner 1766)subelongatus(Motschulsky, 1860)typographus(Linnaeus, 1758)montanus(Eichhoff, 1881)perturbatus(Eichhoff, 1869)(Continued)tridens(Mannerheim, 1852)Monarthrumfasciatum(Say, 1826)mali(Fitch, 1855)Orthotomicuserosus(Wollaston, 1857)caelatus(Eichhoff, 1868)latidens(LeConte, 1874)Pagiocerusfrontalis(Fabricius, 1801)PhloeosinuscupressiHopkins, 1903punctatusLeConte, 1876sequoiaeHopkins, 1903Phloeotribusliminaris(Harris, 1852)scarabaeoides(Bernard, 1788)Pityoboruscomatus(Zimmermann, 1868)Pityogeneschalcographus(Linnaeus,1760)PityophthorusjuglandisBlackman, 1928PolygraphusproximusBlandford, 1894occidentalisSchedl, 1954rufipennis(Kirby, 1837)Pseudipsconcinnus(Mannerheim, 1852)Pseudohylesinusgranulatus(LeConte, 1868)Pseudopityophthorusminutissimus(Zimmermann, 1868)pruinosus(Eichhoff, 1878)Scolytusintricatus(Ratzeberg, 1837)morawitzi(sem*nov, 1902)multistriatus(Marsham, 1802)ratzeburgi(Janson, 1856)SuperfamilyFamilySubfamilyGenusSpeciesAuthor and yearTable 11.3 (Continued)rugulosus(Müller, 1818)schevyreviSemanov, 1902scolytus(Fabricius, 1775)ventralisLeConte, 1868Tomicusdestruens(Wollaston, 1865)minor(Hartig, 1834)piniperda(Linnaeus, 1758)Truncaudumagnatum(Eggers, 1923)Trypodendrondomesticum(Linnaeus, 1758)Xyleborusdispar(Fabricius, 1792)ferrugineus(Fabricius, 1801)fornicatusEichhoff, 1868glabratusEichhoff, 1877perforans(Wollaston, 1857)spp.—costatomorphusSchedl, 1950monographus(Fabricius, 1792)Xylosandruscompactus(Eichhoff, 1876)crassiusculus(Motschulsky, 1866)germanus(Blandford, 1894)morigerus(Blandford, 1894)Xyloterinuspolitus(Say, 1826)Xyloteruslineatus(Olivier, 1795)Subfamilies, genera and species are listed in alphabetical order within each family. All families except Carabidae belong to the suborder Polyphaga; the classification follows Table 11.1.Insect Biodiversity: Science and Society378can expect threats to our food supply and global biodiversity (Allen‐Wardell etal. 1998).11.3.2 Beetles ofCultural ImportanceBeetles have cultural significance to societies around the world. The most ubiquitous of these cultural practices is the use of beetles as a food and the traditions around that use. Beetles are recog­nized by most indigenous societies as good food. Onore (1997) found that at least 30 species of bee­tles were eaten in Ecuador, including many large and common beetles found near settlements in the Ecuadorian highlands. Smith and Paucar (2000) provide a detailed account of the traditions surrounding the eating of the scarab beetle Platycoelia lutescens. Based on these accounts, the practice of entomophagy seems to be widespread and important to Ecuadorian society, especially among the poor. Ecuadorians are generally famil­iar with how and when to collect and prepare these beetles for food (Smith and Paucar 2000). In another example, Utsunomiya and Masumoto (1999) found striking similarities between beetle consumption practices in northern Thailand and Ecuador. In northern Thailand, approximately 100 different species of beetles are eaten, with 89% of respondents listing “taste” as the answer to the question of why insects are eaten. Cultural tradi­tions surrounding beetle consumption are not restricted to Ecuador and Thailand. They also have been observed in many other world cultures. However, these traditions might be in decline because of apparent disdain for eating insects in cultures of European origin, and competition with vertebrate‐based agricultural production.Perhaps the best‐known use of beetles in mythology and religion is the sacred scarab of the ancient Egyptians. The scarab beetle symbol was used prominently over approximately 3000 years, from the First Dynasty (5000 years ago) (Cambefort 1994) to the conquest of Egypt by the Roman Empire. Ancient Egyptians likened the rolling of dung balls by dung beetles to the sun rolling across the sky each day. Because the sun was thought to be reborn each day, scarab beetles became powerful symbols of resurrection and eternal life, which were prominent aspects of ancient Egyptian mythology. Through 3000 years of ancient Egyptian cultural development, an elaborate mythology developed around scarab beetles, including the incorporation of scarab pupation (likened to mummification) and other aspects of their natural history and life cycle into mythological stories. Cambefort (1994) gives a detailed review of all facets of the use of scarab beetles by ancient Egyptians. Kritsky (1991) and Cambefort (1994) reported that the carvings and symbols of Buprestidae and Elateridae also held meaning to the ancient Egyptians. They hypoth­esized that buprestids symbolized the Egyptian myth of rebirth, perhaps due to the emergence of adult beetles from the trunks of living trees.In many countries, keeping personal beetle col­lections and live beetles are popular hobbies, especially in Europe, Japan, and North America. Beetles are a major component of the multimil­lion dollar insect trade. This trade is conducted by mail and Internet or at annual insect fairs in cities including Los Angeles, Paris, Prague, and Tokyo. Keeping live beetles is popular in Japan, where an industry provides not only live beetles, but the required food and paraphernalia for keeping them alive. Evidence for the popularity of this pas­time in Japan can be seen on the website www.youtube.com, which features numerous Japanese videos of live beetles.11.3.3 Beetles ofMedical andLegal Importance11.3.3.1 Medical EntomologyBlistering of human skin (dermatosis) can be caused by beetles of the families Meloidae, Oedemeridae, and Staphylinidae (Paederus and Paederidus) (Frank and Kanamitsu 1987, Nicholls et al. 1990). Dermatosis occurs when a beetle’s hemolymph is released onto the skin after it is crushed or touched accidentally. The vesicant chemical compound in the body fluids,of the Meloidae and Oedemeridae is cantharidin, whereas pederin is the vesicant in beetles of the genera Paederus and Paederidus (Mackie et al. 1945, Nichols etal. 1990, Piel etal. 2005). These 11 Biodiversity ofColeoptera 379chemicals are generally used by the beetles against predators (Pinto and Bologna 2002).Pederin belongs to a group of complex com­pounds found only in these staphylinid genera and a few marine sponges (Piel etal. 2004a). This com­pound is found only in females of some species and is thought to be produced by symbiotic bacte­ria (Kellner 2001). Research on the biosynthesis and mode of action of pederin has increased since the discovery that related compounds have potent antitumor properties (Piel etal. 2004a, b). Pederin inhibits protein biosynthesis in tumor cells, and the isolation of pederin‐producing genes in these symbiotic bacteria might lead tothe development of anticancer drugs (Piel etal. 2005).Canthariasis is a term used to describe the infection of human internal organs by beetles. The most common type of canthariasis occurs when people accidentally or voluntarily ingest beetles. Accidental ingestions generally occur when people eat foods contaminated by stored‐product pests. Adults, larvae, and cast skins can be ingested in this manner, resulting in irrita­tion of the digestive system or allergic responses. The inadvertent ingestion of larder beetle larvae (Dermestes lardarius, Dermestidae), which are covered by long, narrow, and barbed setae, can cause diarrhea, abdominal pain, and perianal itch (Goddard 2000).Deliberate ingestion of beetles for medicinal purposes also has also been reported. Cantharidin is a well‐known insect‐derived medicinal com­pound from the families Meloidae and Oedemeridae. The insect referred to as Spanish fly (Lytta vesicatoria) is a European meloid beetle. Dried, crushed beetles containing cantharidin have been ingested as a vesicant to treat ailments and as an aphrodisiac for millennia (Karras etal. 1996). Cantharidin can be toxic to humans, caus­ing significant illness (Sandorini 2001). Ingestion of live darkling beetles (Ulomoides dermestoides) for similar purposes was reported from Southeast Asia (Sandroni 2001). The practice of ingesting uncooked beetles for food or medicine is risky because some are intermediate hosts for tape­worm and nematode parasites of vertebrates (Mackie etal. 1945, Lethbridge 1971, Halffter and Matthews 1966), possibly including humans (Chu etal. 1977).One type of canthariasis, called scarabiasis, refers to the short‐term infestation of the human gut by adult dung beetles (Scarabaeidae). This condition usually affects preschool children in the tropics. The infestation begins when dung beetles fly into dwellings with sleeping children. Adult dung beetles, usually shorter than 1 cm, follow the smell of feces and are thought to enter the anus and feed internally (Arrow 1931, Halffter and Matthews 1966). Adult beetles are often seen flying away from newly passed stool (Rajapakse 1981). Other types of canthariasis involve the infrequent cutaneous, nasal, ocular, and urinary infections of humans caused by beetle eggs, larvae, or adults (Mackie etal. 1945).Several families of elateroid beetles produce light, using a biochemical reaction (Lloyd 1983, Viviani 2002). Light is produced when enzymes called luciferases catalyze the oxidization of luciferin compounds. Beetles principally use this cold light, called bioluminescence, for sex­ual communication, although other possible functions, including aposematic signals, attrac­tion of prey, and defense, are possible (Lloyd 1983, Underwood etal. 1997). Studies of firefly (Lampyridae) light production have resulted in the development of several medical applica­tions, based on the use of luciferases and their associated genes. These applications range from monitoring the progress of infections such as HIV (Contag et al. 1997) to visualizing living cells in human embryonic development (Greer and Szalay 2002). Luciferases have played a sig­nificant role in the development of more effi­cient drugs for many diseases (Viviani 2002).11.3.3.2 Forensic EntomologyThe major contribution of insects in criminal cases involving homicide is to estimate the lim­its of the postmortem interval (time between death and discovery of the body). Flies are dom­inant on human corpses, although several groups of beetles are also often present, either preying on other arthropods or feeding on the body itself (Smith 1986, Catts and Haskell 1990). Insect Biodiversity: Science and Society380The postmortem interval is usually estimated from experimental studies using pigs or other vertebrates, and then applied to human corpses in criminal cases (Tabor et al. 2004, Arnaldos etal. 2005). The period of development of the beetles and the succession of species that colo­nize carcasses in various stages of decomposi­tion provide data for estimating the postmortem interval (Franc et al. 1989, Moura et al. 1997, Carvalho etal. 2000, Turchetto and Vanin 2004). The succession of beetle species tends to follow a rough pattern from an initial fresh stage of decomposition through bloated and active decay stages to a final dry stage. The Dermestidae (skin beetles) and Cleridae (bone beetles) are among the most common beetles on corpses and have provided important postmortem information, especially for finds of dry skeletal remains (de Souza and Linhares 1997, Kulshresta and Satpathy 2001).Although most experimental postmortem‐interval studies use bodies on the ground surface, some research has examined buried or sub­merged bodies (Smith 1986). A few beetles (e.g., Histeridae, Silphidae, and Staphylinidae) are associated with buried bodies (Payne etal. 1968, VanLaerhoven and Anderson 1999, Bourel etal. 2004). Molecular and toxological analyses show that necrophagous beetles also could be informa­tive for criminal cases involving badly decom­posed bodies (DiZinno etal. 2002), movement of bodies (Benecke 1998), and bodies with toxic substances (e.g., drugs, heavy metals, and poi­sons) (Bourel etal. 2001, Gagliano‐Candela and Aventaggiato 2001).11.3.4 Beetles asResearch ToolsBeetles are used widely as research tools in bio­physics and related disciplines. Because they are the most diverse animal order, beetles possess great potential for bioengineering studies. Geometry and mechanics of elytral opening and closing are studied by aeronautic and astronautic engineers (Frantsevich etal. 2005). Whirligig bee­tles (Gyrinidae), roughly 1 cm in length, swim on the water surface at 55 cm per second and are capable of making 12 horizontal rotations per sec­ond (Fish and Nicastro 2003), which would be a dream performance for any human‐made autono­mous device. Aquatic larvae of Hydrobius fuscipes (Hydrophilidae) demonstrate how simple side‐to‐side body swimming movements allow a novel kind of limbless skating on the lower surface of the water, with tracheal gills acting as anchors (Brackenbury 1999). Walking mechanics in Pachnoda marginata (Scarabaeidae) involve fric­tion forces between the tarsal claw systems and walking substrates (Dai et al. 2002). Specialized insect adhesive devices, such as the arolium, euplantulae, pulvilli, and tarsal hairs (Gorb 2001), inspire engineers to develop novel adhesive sur­faces. Legs of the leaf beetle Gratiana spadicea (Chrysomelidae) attach to leaf surfaces by match­ing tarsal claw aperture with that of pointed rays of the host‐plant trichomes (Medeiros and Moreira 2002). Wide, bilobed tarsi of rove beetles in the genus Stenus (Staphylinidae) allow the beetles to run well on solid surfaces or water (Betz 2002).The high diversity of beetles makes them use­ful tools for physiological research. Studies of muscle function in arthropods, using the beetle Cotinis mutabilis (Scarabaeidae),,suggest that asynchronous flight muscles can provide greater power output than can synchronous muscles for operation at the high‐contraction frequencies of insect flight (Josephson etal. 2001). Some dung‐rolling scarab beetles (Scarabaeus species) pos­sess a strong sensitivity to polarized light, a sense lacking in humans. The crepuscular beetle Scarabaeus zambesianus rolls dung balls away from the dung pile to avoid competition, navigat­ing by polarized skylight sensed by specialized ommatidia of the dorsal rims of its eyes (Dacke etal. 2003). Biophysical and behavioral studies of bioluminescent insects, including click beetles (Elateridae) and fireflies (Lampyridae), provide insights into the evolution of color vision (Stolz 2003, Booth etal. 2004). Discoveries of the abili­ties of tiger and scarab beetles (Carabidae: Cicindelinae and Scarabaeidae) to use paired membranous ears to detect airborne sounds 11 Biodiversity ofColeoptera 381(Forrest etal. 1997) shed light on the evolution of this sense in the Animalia.Many phytophagous beetles rely on aggrega­tion pheromones to coordinate behaviors. Bark beetles use pheromones to coordinate attacks on host trees (Raffa and Berryman 1983). The dis­covery of the first pheromone for the Colorado potato beetle, Leptinotarsa decemlineata, is unu­sual because it is the first male‐produced phero­mone known in the Chrysomelidae (Dickens etal. 2002). In longhorn beetles (Cerambycidae), males use sex pheromones to attract females or aggre­gate (Lacey etal. 2004).Beetles are particularly important research tools for biochemistry and DNA research. Tribolium castaneum, the red flour beetle (Tenebrionidae), commonly consumes stored grains worldwide. Like the famous fruit fly, Drosophila melanogaster, it is used widely in genetics and developmental biology research, and was the first beetle species whose complete genome sequence was published. Its genome consists of about 200 million nucleotides arranged in a haploid set of 10 chromosomes. For comparison, the human genome has about 15 times more nucleotides and a haploid set of 23 chromosomes. Tribolium castaneum was the first animal reported to produce inhibitors of prostaglandin synthetase, which were purified from the beetle’s defensive secretions (Howard etal. 1986). These substances are used widely in aspirin‐like anti‐inflammatory drugs. This bee­tle is also intensively studied as a model for understanding the mechanisms of insect resist­ance to insecticides.Using beetles as research tools sheds light on bizarre aspects of evolution, some of which are unique. For example, Micromalthus debilis, the sole North American member of the family Micromalthidae, possesses one of the most bizarre life cycles in the Metazoa. This species combines both thelytokous and viviparous larviform diploid females and rare haploid males, which eat and kill their own mothers from inside (Pollock and Normark 2002). The second known case of hap­lodiploid sex determination in beetles is that of some scolytine weevils including Ozopemon from Southeast Asia, which is also the only beetle genus with neotenic and strangely modified larviform males (once thought to be Histeridae) (Jordal etal. 2002). The huge diversity of ecological relation­ships with other organisms makes beetles ideal for researching the origins of sociality (Scott 1998), parasitism (Weber etal. 2006), symbiosis (Kellner 2003), and phoresy (Bologna and Pinto 2001).Beetle groups with well‐resolved phylogenies also allow biogeographical and paleogeographical reconstructions. Most commonly, these recon­structions are achieved using DNA data for spe­cies with weak dispersal capacities, such as flightless Scarabaeus dung beetles (Scarabaeidae), which show grades of colonization of the Namib Desert during the Miocene (Sole et al. 2005). Phylogenetic trees inferred from Nesotes darkling beetle (Tenebrionidae) mitochondrial DNA (mtDNA) suggested how the genus colonized the Canary Islands (Rees et al. 2001). The well‐resolved phylogeny of endemic Iberian diving beetles (Dytiscidae) indicated that their specia­tion was induced by repeated fragmentation of populations during glacial and interglacial peri­ods (Ribera and Vogler 2004). Patterns of insect colonization of Pacific islands were deduced based on the distribution and phylogeny of Colymbetinae diving beetles (Dytiscidae) on New Caledonia and Fiji (Balke etal. 2007). Analysis of fossil and subfossil chitinous remains of beetles in Quaternary sediments (1.8 mya–present) pro­vides quantitative estimates of ancient tempera­tures and precipitation levels (Porch and Elias 2000). All these studies capitalized on the diver­sity and relative abundance of beetles to focus on more inclusive natural phenomena.Fireflies (Lampyridae) are central to biolu­minescence research (McElroy and DeLuca 1983, Viviani 2002). In addition to its medical applications, bioluminescence research has yielded several other commercial applications. For example, luciferases are routinely used as environmental biosensors to monitor levels of pollutants such as agrochemicals, lead, and mercury (Naylor 1999).Insect Biodiversity: Science and Society38211.3.4.1 DNA Barcoding ofBeetles–ANorth American Case StudyBeetles represent one of the taxonomically bet­ter‐known major orders of North American insects. Diversity estimates are available from Bousquet et al. (2013) for Canada and from Arnett and Thomas (2000), Arnett etal. (2002), and Marske and Ivie (2003) for North America as a whole. A concerted effort was recently initiated to document the diversity of beetles in North America using DNA barcodes. Here, we examine its progress, expand on some findings, and dis­cuss remaining gaps in barcoding coverage.The Canadian National Collection of Insects, Arachnids, and Nematodes (CNC; Ottawa, Ontario, Canada) has recently (2010–14) collabo­rated with the International Barcode of Life Project (University of Guelph, Ontario, Canada) to build a barcode‐sequence library for the beetles of Canada. Secondarily, many beetle taxa have been barcoded from the United States to increase North American coverage. This effort was central to increasing the number of North American bee­tle species with high‐quality barcode data (> 500 base pairs of the cytochrome oxidase subunit I gene (COI)) in the Barcode of Life Data System (BOLD: http://www.boldsystems.org) from 1613 to 6905.The barcoding diversity and coverage were examined using the barcoding index number (BIN) system in BOLD detailed by Ratnasingham and Hebert (2013). BINs can approximate spe­cies diversity well, but are discordant with cur­rent taxonomy in roughly 10% of cases (Ratnasingham and Hebert 2013). According to Ratnasingham and Hebert (2013), these limita­tions are due to four main factors: taxonomic error, sequence contamination, problems with the BIN calculation methodology, and lack of COI‐sequence variation due to introgression or their evolutionarily young age. In our observa­tions, misidentification of specimens is also a confounding factor when working with the BOLD data, as samples come from multiple sources and are not all identified by experts. For this study, barcodes were only considered if they met the 500 base‐pair threshold for quality, fol­lowing Ratnasingham and Hebert (2013).The barcoding coverage for the beetles of Canada and North America is summarized in Table 11.4, Fig. 11.3, and Fig. 11.4. The number of BINs divided by the number of known spe­cies approximates barcoding coverage well, with a few caveats. First, the number of known spe­cies is only an approximation for most North American families, as there are still many gaps in basic taxonomic knowledge of these groups. The number of known species is less than the actual number of species for most families because many North American,families have not undergone thorough taxonomic revisions and many beetle species remain undescribed. In addition, the number of BINs contains a degree of error for the reasons mentioned above, and overestimates or underestimates the actual number of species sampled roughly 10% of the time. Beyond this, barcoding sampling of North American beetles is still in its early stages and lacks coverage of much of the observed varia­tion in described species.Our results show that 49.7% of described Canadian beetle species and 27.4% of North American beetle species have been barcoded in BOLD. The percentage is significantly higher for Canada, reflecting the specific efforts there and the lower diversity at northern latitudes. Our general observations of the coverage are what one might expect: easily collected taxa (abundant, easily observed, and with a wide dis­tribution) are well represented, whereas diffi­cult to collect taxa (rare, cryptic habits, and with limited distribution) are poorly repre­sented. The former is exemplified by the Silphidae and Geotrupidae (76.7% and 78.6% North American coverage, respectively), as these families are relatively well known and have low diversity in North America, and most spe­cies are fairly widespread, readily observed and collected, and easily attracted to traps. The lat­ter is exemplified by the Histeridae and Ripiphoridae (3.9% and 5.9% North American coverage, respectively), as these families are also reasonably well known taxonomically but have Table 11.4 Estimated number ofColeoptera species forNorth America andCanada, withtheassociated number ofDNA‐barcoding BINs andthecalculated percent regional barcoding coverage.SuborderSuperfamilyFamilyNorth America speciesNorth America BINsNorth America percentage coverageCanada speciesCanada BINsCanada percentage coverageArchostemata—Cupedidae44100.0033100.00Micromalthidae11100.00100.00MyxophagaSphaeriusoideaHydroscaphidae100.00000.00Sphaeriusidae3133.33000.00Adephaga—Gyrinidae563358.93342573.53Trachypachidae4250.0022100.00Rhysodidae8337.52150.00Carabidae2,40292938.6898958158.75Haliplidae671522.39341029.41Noteridae1417.142150.00Amphizoidae3266.673266.67Dytiscidae51319638.2128416257.04PolyphagaHydrophiloideaHydrophilidae25812448.061517952.32 Sphaeritidae100.00100.00Histeridae435173.9113685.88StaphylinoideaHydraenidae6757.4627414.81Ptiliidae1173025.64492959.18Agyrtidae11218.188112.50Leiodidae3248726.851827139.01Silphidae302376.67261869.23Staphylinidae4,37093921.491,68273643.76ScarabaeoideaPleocomidae2700.00000.00Geotrupidae282278.5712866.67Passalidae4125.00100.00(Continued)SuborderSuperfamilyFamilyNorth America speciesNorth America BINsNorth America percentage coverageCanada speciesCanada BINsCanada percentage coverageTrogidae431125.5815426.67Glaresidae25416.00200.00Diphyllostomatidae300.00000.00Lucanidae261038.4614857.14Ochodaeidae35617.144125.00Hybosoridae5480.0011100.00Glaphyridae8112.50100.00Scarabaeidae1,70058634.4722114364.71ScirtoideaEucinetidae11654.557685.71Clambidae12325.007342.86Scirtidae503468.002531124.00DascilloideaDascillidae500.00000.00Rhipiceridae5120.00100.00BuprestoideaSchizopodidae700.00000.00Buprestidae76217122.441779151.41ByrrhoideaByrrhidae352160.00261869.23Elmidae992525.2532721.88Dryopidae1300.00600.00Lutrochidae300.00100.00Limnichidae2800.00300.00Heteroceridae341647.0628828.57Psephenidae16637.504250.00Ptilodactylidae191052.6346150.00Chelonariidae100.00000.00Eulichadidae100.00000.00Table 11.4 (Continued)SuborderSuperfamilyFamilyNorth America speciesNorth America BINsNorth America percentage coverageCanada speciesCanada BINsCanada percentage coverageCallirhipidae100.00000.00ElateroideaArtematopodidae88100.0055100.00Brachypsectridae100.00000.00Cerophytidae200.00000.00Eucnemidae851720.00381436.84Throscidae2021105.00817212.50Elateridae96543645.1838623861.66Lycidae763748.68292482.76Telegeusidae300.00000.00Phengodidae23939.1312200.00Lampyridae1244838.71322475.00Omethidae1000.00000.00Cantharidae47311624.521549259.74DerodontoideaDerodontidae9555.568450.00Nosodendridae400.00200.00Jacobsoniidae100.00000.00BostrichoideaDermestidae1172924.79492448.98Endecatomidae200.00100.00Bostrichidae752026.6724729.17Ptinidae4714810.19983131.63LymexyloideaLymexylidae2150.0011100.00CleroideaTrogossitidae591118.642229.09Cleridae2434418.11522242.31Melyridae5208516.35532343.40CucujoideaByturidae2150.0011100.00Sphindidae9555.566466.67Biphyllidae300.00100.00Erotylidae822834.15281450.00Monotomidae551120.0027829.63(Continued)Cryptophagidae1455034.48734663.01Silvanidae32618.7516531.25Cucujidae8562.508562.50Passandridae300.00100.00Phalacridae1221411.48810125.00Laemophloeidae5259.6213323.08Kateretidae11436.368337.50Nitidulidae1737241.621015756.44Smicripidae200.00000.00Bothrideridae1800.00400.00Cerylonidae19315.798337.50Endomychidae45817.7816637.50Coccinellidae48114730.5616110967.70Corylophidae611321.31161381.25Latridiidae1406747.86645890.63TenebrionoideaMycetophagidae261142.3116850.00Archeocrypticidae100.00000.00Ciidae841517,.86291034.48Tetratomidae26726.9220630.00Melandryidae502754.00432660.47Mordellidae1899650.79755978.67Ripiphoridae5135.8811218.18Zopheridae361130.5619421.05Tenebrionidae1,18435029.561416948.94Table 11.4 (Continued)SuborderSuperfamilyFamilyNorth America speciesNorth America BINsNorth America percentage coverageCanada speciesCanada BINsCanada percentage coverageProstomidae100.00100.00Synchroidae22100.0022100.00Stenotrachelidae10660.009555.56Oedemeridae8766.9013430.77Meloidae4245512.97472553.19Mycteridae1218.334125.00Boridae2150.002150.00Pythidae77100.0066100.00Pyrochroidae501326.00221359.09Salpingidae20525.0015426.67Anthicidae2296126.64652741.54Aderidae4824.1710110.00Scraptiidae493979.592236163.64ChrysomeloideaCerambycidae95830131.4236819853.80Megalopodidae9333.337342.86Orsodacnidae4125.0011100.00Chrysomelidae1,86948726.0659826644.48CurculionoideaNemonychidae15853.338675.00Anthribidae882225.00201365.00Belidae200.00000.00Attelabidae512549.02141392.86Brentidae1528757.244950102.04Dryophthoridae841720.2427725.93Brachyceridae412560.98191157.89Curculionidae2,79447416.9682327433.29Total25,1066,89427.468,2374,09649.73Numbers originate from Arnett and Thomas (2000), Arnett etal. (2002), and Marske and Ivie (2003) for North American species; Bousquet etal. (2013) for Canadian species; and the Barcode of Life Data System (BOLD: http://www.boldsystems.org) for North American and Canadian barcoding index numbers (BINs).Insect Biodiversity: Science and Society388ThroscidaePhengodidaeScraptiidaePtilodactylidaeLeiodidae, Endomychidae, Cerylonidae, Kateretidae, Eucnemidae, Ciidae, Curculionidae, Ptinidae, Silvanidae, OedemeridaePhalacridae, ScirtidaeBrentidaePythidae, Artematopodidae, Cupedidae, Trachypachidae, Synchroidae, Orsodacnidae, Byturidae, Lymexylidae, Hybosoridae, Attelabidae, LatridiidaeEucinetidae, Lycidae, CorylophidaeMordellidae, Lampyridae, Nemonychidae, GyrinidaeSilphidae, Coccinellidae, Geotrupidae, Sphindidae, Amphizoidae, Anthribidae, Scarabaeidae, Cryptophagidae, Cucujidae, Elateridae, MelandryidaeCantharidae, Ptiliidae, Pyrochroidae, Carabidae, Brachyceridae, Lucanidae, Dytiscidae, Nitidulidae, Stenotrachelidae, Cerambycidae, Meloidae, Hydrophilidae, BuprestidaeErotylidae, Mycetophagidae, Derodontidae, Psephenidae, Noteridae, Rhysodidae, Boridae, Dermestidae, Tenebrionidae, Chrysomelidae, Staphylinidae, Melyridae, Clambidae, Megalopodidae, Cleridae, AnthicidaeRipiphoridae, Hydraenidae, AgyrtidaeTetratomidae, Monotomidae, Haliplidae, Bostrichidae, Heteroceridae, Salpingidae, Trogidae, Dryophthoridae, Mycteridae, Ochodaeidae, Laemophloeidae, Elmidae, ZopheridaeAderidae, Trogossitidae, HisteridaeDryopidae, Bothrideridae, Limnichidae, Nosodendridae,Ischaliidae, Glaresidae, Lutrochidae, Biphyllidae, Passandridae, Endecatomidae, Prostomidae, Sphaeritidae, Glaphyridae, Rhipiceridae, Passalidae, Micromalthidae0 50 100 150 200Figure 11.3 Percentage (aggregated in 10% increments) of Canadian Coleoptera species included in the Barcode of Life Data System (BOLD: http://www.boldsystems.org). Percentage coverage is the number of barcoding index numbers (BINs) for a family divided by the number of known species (from Table 11.4). Percentage coverage values of more than 100% indicate poorly known groups in need of further taxonomic and survey research.11 Biodiversity ofColeoptera 389ThroscidaeArtematopodidae, Pythidae, Cupedidae,Synchroidae, MicromalthidaeScraptiidaeGeotrupidae, Silphidae, HybosoridaeTenebrionidae, Elmidae, Trogidae, Ptiliidae, Pyrochroidae, Chrysomelidae, Anthicidae, Bostrichidae, Leiodidae, Tetratomidae Scirtidae, AmphizoidaeCucujidae, BrachyceridaeGyrinidae, Derodontidae, Sphindidae, Brentidae, Byrrhidae, StenotrachelidaePtilodactylidae, Nemonychidae, Melandryidae, Eucinetidae, MordellidaeLatridiidae, Hydrophilidae, Lycidae, Attelabidae, Elateridae, Heteroceridae, Trachypachidae, Boridae, Byturidae, Lymexylidae Nitidulidae, MycetophagidaePsephenidae, Rhysodidae, Dytiscidae, Lucanidae, Carabidae, Lampyridae, Phengodidae, KateretidaeMegalopodidae, Sphaeriusidae, Erotylidae, Scarabaeidae, Cryptophagidae, Zopheridae, Coccinellidae, CerambycidaeEndomychidae, Ciidae, Cleridae, Agyrtidae, Trogossitidae, Silvanidae, Cerylonidae, Glaresidae, Melyridae, Curculionidae, Ochodaeidae, Eucnemidae, Monotomidae, RhipiceridaeCantharidae, Dermestidae, Dryophthoridae, Corylophidae, Staphylinidae, Haliplidae, Buprestidae, Anthribidae, Salpingidae, Clambidae, Orsodacnidae, PassalidaeGlaphyridae, Meloidae, Ptinidae, PhalacridaeMycteridae, Laemophloeidae, Ripiphoridae, Oedemeridae, Noteridae, HydraenidaeHisteridae, AderidaeLimnichidae, Pleocomidae, Bothrideridae, Dryopidae, Omethidae, Schizopodidae, Dascillidae, Nosodendridae, Lutrochidae, Biphyllidae, Diphyllostomatidae, Ischaliidae, Passandridae, Telegeusidae,Belidae, Cerophytidae, Endecatomidae, Smicripidae, Archeocrypticidae, Brachypsectridae, Callirhipidae, Chelonariidae, Eulichadidae, Hydroscaphidae, Prostomidae, Sphaeritidae, Jacobsoniidae0 25 50 75 100Figure 11.4 Percentage (aggregated in 5% increments) of North American Coleoptera species included in the Barcode of Life Data System (BOLD: http://www.boldsystems.org). Percentage coverage is the number of barcoding index numbers (BINs) for a family divided by the number of known species (from Table 11.4). Percentage coverage values of more than 100% indicate poorly known groups in need of further taxonomic and survey research.Insect Biodiversity: Science and Society390many species with cryptic habits; they either burrow in substrates or are myrmecophiles (Histeridae), termitophiles (Histeridae), or par­asitoids (Ripiphoridae). As such, many species in these families have short activity periods as adults and are difficult to collect without taxon‐specific techniques. Other examples of taxa with no barcode coverage are the families Pleocomidae and Diphyllostomatidae, both of which are endemic to the West Coast of North America, with small distributions, and are not generally found unless collectors specifically seek them. Of the 129 North American beetle families, 27 (21%) are not yet represented with high‐quality barcoding data in BOLD, but most of these families are known only from one to three species with restricted distributions.For the Canadian fauna, there was an early initiative from 2006 to 2010 to do a barcode inventory of the beetles of Churchill, Manitoba (Woodco*ck et al. 2013). The resulting data set was augmented by the Biodiversity Institute of Ontario through intensive sampling around Guelph, Ontario, Canada, and additional sam­pling mainly in national parks across Canada. In 2011, a joint effort between the Biodiversity Institute of Ontario and the CNC was initiated to,barcode Canadian beetle taxa using pinned specimens in the collection. A first run through all taxa was completed in 2014, with additional sampling still ongoing. The results of these combined efforts yielded 50% barcoding cover­age of Canadian beetle species. This level of coverage is higher than for all of North America to the point where most common or widespread species are represented by barcodes. Barcode coverage of 50% of described species or more has been achieved for 57 of the 112 families of Canadian beetles, including diverse families such as the Carabidae, Cerambycidae, and Elateridae. Significant gaps in coverage remain for three of the most diverse families: Chrysomelidae, Curculionidae, and Staphylinidae, which have less than 50% bar­code coverage for Canadian species. The gaps in coverage could be efficiently addressed by tar­geting these groups for future work.Some families are so poorly known that general barcode efforts have already found many more BINs than the number of described species. This is so for the families Scirtidae, Scraptiidae, and Throscidae, which have 105.0%, 84.8%, and 68.0% barcoding coverage for North America and 212.5%, 180.0%, and 124.0% for Canada. It is obvi­ous from examining the barcoding data that many more species are present in the focal regions than are indicated by the number of described species. These families and others are fertile ground for taxonomists, and highlight the poor state of tax­onomy in the backyards of some of the top ento­mology museum collections in the world.Two North American genera, Phyllophaga and Serica (Scarabaeidae: Melolonthinae), were examined for BIN discordance. They were selected because of their high diversity (212 and 100 North American species, respectively; Evans and Smith 2009), with low variation in external morphology. These two factors make these gen­era likely to have poor COI lineage sorting between species because their similar morphol­ogies suggest that their diversity is a result of a recent speciation explosion. The identification of species in these genera can be accurate using the highly diagnostic genitalic characters. Among BOLD specimens, 16 of the 143 Phyllophaga BINs and four of the 36 Serica BINs were tagged as discordant, with more than one species based on morphology included (both approximately 11%). This percentage is in line with the findings of Ratnasingham and Hebert (2013) and probably represents the high end of expected BIN discordance error for a group with high recent speciation and poor lineage sorting (whereby descendants of an ancestor species inherit different subsets of variants of the ances­tral mitochondrial genome). A good case study for Coleoptera would be to determine whether this percentage changes when more species and more specimens within species are sampled.DNA Barcoding Detects New Invasive Species Any inventory work can detect new taxa for a focal region, including new species, range exten­sions, or invasive species. Our analysis of the 11 Biodiversity ofColeoptera 391barcoding data for North American Coleoptera detected a new invasive species in Canada.Anthribus nebulosus (Anthribidae) is a preda­tor of scale insects (Hemiptera: Sternorrhyncha), that was intentionally introduced to Virginia, United States, in 1978–79. It is thought to have been introduced elsewhere in North America because of its disjunct distribution (Hoebeke and Wheeler 1991). This Eurasian species also has been recorded from southern New England (Hoebeke and Wheeler 1991) and adjacent regions of Pennsylvania and New Jersey (http://bugguide.net/node/view/278376/data). The bar­coding records from in and around Guelph, Ontario, Canada, were unexpected because this species had not been observed to spread rap­idly and was not first detected in Canada near known localities in the United States. It is unclear whether this disjunct distributional record is a result of a local introduction or an unde­tected general expansion of the species range. Specimens of A. nebulosus were collected in and near Guelph in three separate collecting events and barcoded as part of general survey and inventory efforts. The barcoding data were included in overall Coleoptera and Curculionoidea analyses and were a match within a BIN including identified specimens from Germany. Robert Anderson (Canadian Museum of Nature, Ottawa, Ontario, Canada) confirmed the identification of Guelph speci­mens. Additional specimens from Ontario in the same BIN were collected in Rouge River National Urban Park and on Beausoleil Island (collectively new country and provincial records).A false‐positive record for an invasive species resulted from a BIN from the root weevil genus Polydrusus (Curculionidae: Entiminae), which is widespread globally. Four native and three invasive species are known from North America (Anderson 2002). Specimens from Point Pelee, Guelph, and Rouge River, Ontario, Canada, were collected as part of a general survey to build the barcode library. Our analysis of bar­coding data revealed that these specimens shared a BIN with German specimens identi­fied as Polydrusus corruscus. This is a Eurasian willow‐feeding species (Salix), not previously known from North America. The Ontario specimens deposited in the Biodiversity Institute of Ontario were examined and identi­fied as Polydrusus impressifrons, an invasive species previously recorded from North America. The error seems to have originated from misidentified German specimens. This case shows the need to verify identifications of specimens in BOLD before using its results to infer new records of invasive species or other discoveries.DNA Barcoding Reexamines Species Limits of Holarctic Taxa Species with a Holarctic distribution are fairly common. Depending on their range and dispersal histories, Nearctic and Palearctic pop­ulations of apparent Holarctic species might have been isolated long enough for lineage sort­ing to occur in fast‐evolving gene regions and for the two populations to diverge enough to be considered separate species. Barcoding can be a rapid way to test for such cryptic species. When the Palearctic and Nearctic specimens are sorted to separate BINs, these species should be studied further to determine whether they are different species, using an integrative taxonomic approach. If the barcoding results are supported by morphological, host plant, behavioral, or other characters, then broad‐based support has been achieved and the two species should be considered distinct and valid.The barcoding survey and inventory work in North America and Eurasia has examined numerous Holarctic beetle species. Barcoding has detected two separate BINs (one North American and the other Eurasian) for some spe­cies that were formerly considered Holarctic. Species in this category include Ampedus nigri-nus (Elateridae), Dictyoptera aurora (Lycidae), Grypus equiseti (Brachyceridae), Tournotaris bimaculata (Brachyceridae), and Dryocoetes autographus (Curculionidae: Scolytinae). These species should be further investigated to deter­mine whether other data sets support splitting them into separate North American and Eurasian species.Insect Biodiversity: Science and Society392Looking at one specific example, Wood (1982) commented that with D. autographus, “The American specimens tend to have the frons more sparsely granulate, the strial punctures slightly larger and not as deep, and the elytral declivity slightly more convex than the European material. The differences are slight, variable, and not suitable for statistical analysis; there­fore, I prefer to follow Bright (1963) and recog­nize only one species.” These recognized morphological differences coupled with the barcoding results suggest a possible basis for considering the Palearctic and Nearctic popula­tions as two separate species.,Also detected were two examples of species with three different BINs for populations in Eurasia, Alaska, and Canada: Berninelsonius hyperboreus (Elateridae) and Eutrichapion viciae (Brentidae). These examples could indi­cate multiple cryptic species, or perhaps a slow postglacial range expansion with subsequent population isolation and DNA lineage sorting. More evidence is required to determine the number of species present.Adventive beetle species in North America also were analyzed, and some had separate BINs for their North American and Eurasian popula­tions. These species include Brachypterus urti-cae (Kateretidae), Anthocomus equestris (Melyridae), Ischnopterapion loti (Brentidae), and Tanysphyrus lemnae (Brachyceridae). More research is needed to explain these genetic dis­tinctions. Possible explanations include misi­dentifications (as described for Polydrusus), and the possibility that the source populations include multiple cryptic species (e.g., Rhyssemus and Aphodius (Scarabaeidae). It is also possible that the adventive population originated from a source population with an as‐yet‐undetected haplotype (i.e., BIN).DNA Barcoding is Part of the Integrative Taxonomic Approach to Delimiting Species Barcoding and DNA sequence analysis have become a key component of the integrative taxonomic approach to delimiting species. One example is the recent recognition of the scarabs Rhyssemus germanus and Rhyssemus puncticollis as separate species, both occurring in Europe and the latter also occurring in North America. Rhyssemus germanus was described from Germany, and Brown described R. puncticollis much later from Canada. In his description of R. puncticollis, Brown (1929) did not mention R. germanus or any other Palearctic species, imply­ing that this species was endemic to the Nearctic region. Brown (1950) later synonymized his own species under R. germanus, stating that “this species has not been reported previously from America under the name germanus, but comparison of the types of puncticollis with European specimens shows the names to be synonymous.” The assumption that R. germanus was an invasive species to North America, with R. puncticollis as a synonym, was supported by Gordon and Cartwright (1980) and others until recently.Rößner (2012) and Kral and Rakovič (2012) recognized consistent morphological differ­ences between R. germanus and R. puncticollis and revalidated the latter as a valid species, which they record from Germany and the Czech Republic. Barcoding data corroborate the reval­idation of R. puncticollis, showing separate BINs for R. germanus (German specimens) and R. puncticollis (German and Ontario specimens). Independent confirmation robustly supports the hypothesis of two separate and valid species. This new evidence allows researchers to assess whether the true R. germanus occurs in North America and whether R. puncticollis is actually invasive to North America. Four synonyms based on European specimens need to be reex­amined to determine whether they apply to R. germanus or R. puncticollis.Another case involves the cryptic dung‐feed­ing scarab species Aphodius fimetarius and Aphodius pedellus. Until recently, these two species were together named A. fimetarius, thought to be native to Europe and invasive to North America. Wilson (2001) found two chro­mosomal karyotypes in the specimens from Europe and presented subtle morphological characters to support the hypothesis of two 11 Biodiversity ofColeoptera 393cryptic species. This work was supported and expanded by Miraldo etal. (2014), who included specimens from across the entire range, exam­ined COI mtDNA data, and made an in‐depth morphological analysis along with the karyo­typic analysis.Barcoding data from BOLD corroborate the findings of Miraldo etal. (2014), with two clear BINs for A. fimetarius from Europe and California and A. pedellus from Europe and Canada. Aphodius fimetarius and A. pedellus are now understood to be cryptic species; both species occur in Europe and North America and both may be invasive to North America. This example highlights how integrative taxonomy and barcoding can reveal cryptic species among common, widespread taxa.DNA Barcoding Identifies Taxonomic Gaps in Groups with Underestimated Biodiversity The families Scirtidae, Scraptiidae, and Throscidae were iden­tified above as groups where barcoding exposed much hidden species richness. At the generic level, a few spectacular examples are Ptiliolum (Ptiliidae), Cyphon (Scirtidae), Trixagus (Throscidae), and Anaspis (Scraptiidae). Ptiliolum is known from five species in Canada (Bousquet et al. 2013), but pre‐barcoding estimates found “numerous undescribed species occurring in northern and western United States and Canada” (Hall 2000). We found five BINs of Ptiliolum from Alberta and Saskatchewan, Canada, although only one species is known from Alberta and none from Saskatchewan (Bousquet et al. 2013). As Hall (2000) indicated, this genus requires taxo­nomic study because there are many undescribed North American species.The genus Cyphon is also in need of taxo­nomic revision. Young (2002) commented that the genitalic characters are diagnostic but that the external morphology is hom*ogenous among species so that every specimen requires dissec­tion to confirm identifications. This situation has probably led to the poor taxonomy of this group. The barcoding data yield 21 BINs from North America, and Young (2002) mentions 27 described species. The geographic distribution of the described species disagrees with that of the BINs, and there are many more BINs than there are described species in Canada, indicat­ing many undescribed North American species.The genus Trixagus was last revised by Yensen (1975) and has six known species in North America, according to Johnson (2002), who cautioned that “this family is in need of general study at all levels.” Using the barcoding data, we found 13 clear North American Trixagus BINs. This genus is at least twice as diverse in North America as the taxonomic literature indicates. Because most of the barcoding samples were taken from Canada, we can anticipate more undetected BINs and species in the generally more diverse United States. Yensen (1975) com­mented that there was much variation in his species definitions, suggesting several cryptic species, as do the barcoding data. The North American Throscidae need a complete taxo­nomic overhaul. Even with the moderate and general focus of the barcoding to date, the num­ber of North American throscid BINs in BOLD already exceeds the number of known species. This family would be an excellent focal taxon for any taxonomist interested in describing many new North American species.Anaspis has 13 known North American spe­cies but has never undergone a taxonomic revi­sion, and authors have disagreed on whether the genus should be divided into four genera (Pollock 2002). Our barcoding data have recov­ered 13 BINs from North America, but few specimens from these BINs match any of the described species; they also differ in distribu­tion from the described species. Thus, several North American species in this genus remain undescribed.Overall, building the barcode library for North American Coleoptera has reached a point where the data are useful in many ways. Most common or widespread species have been barcoded, so clear matches are likely for any new, unidentified specimens compared with BOLD. The barcoding data also help users detect invasive species, re‐examine species delimitations, identify cryptic species through Insect Biodiversity: Science and Society394integrative taxonomy, and expose gaps in taxo­nomic knowledge. The barcode library remains far from complete, so the next steps are to con­tinue to identify and target taxa with poor cov­erage. This challenge to find,DNA‐quality specimens will increase, as the bulk of the remaining taxa are cryptic species with small geographical ranges.11.4 Threatened BeetlesMany beetles are vulnerable to local and global extinction. These beetles often have low powers of dispersal (flightless), occur only in specific microhabitats, and are distributed over small geographical areas. Beetles of oceanic islands, isolated dunes, caves, mountains, and other eco­logical islands fit into this category. Currently, 791 species appear on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species, of which 12 are listed as extinct, 17 as critically endangered, 47 as endan­gered, and 45 as vulnerable (Table 11.5). Species on the Red List belong to 22 families. Habitat destruction and the introduction of invasive spe­cies continue to threaten most natural ecosys­tems and their myriad beetle species (Spence and Spence 1988, Kamoun 1996, Martikainen and Kouki 2003, Munks etal. 2004, Abellan etal. Table 11.5 Number ofColeoptera species ontheIUCN (2015) Red List ofThreatened Species, by family.Suborder Superfamily Family EX CR EN VU NT LR/NT LC DD TotalAdephaga — Carabidae 1 2 2 2 — 1 — — 8Dytiscidae 6 2 8 6 — — — — 22Polyphaga Staphylinoidea Leiodidae — — — 1 1 — — — 2Silphidae — 1 — — — — — — 1Scarabaeoidea Lucanidae — 4 6 4 1 1 4 — 20Scarabaeidae — 2 15 15 20 — 302 232 586Buprestoidea Buprestidae — — 1 — — — — — 1Byrrhoidea Elmidae — — — 1 — — — — 1Elateroidea Eucnemidae — — — 1 3 — 7 4 15Elateridae — — 3 2 7 — 6 38 56Bostrichoidea Bostrichidae — — — — 1 — 2 — 3Ptinidae — — — — 1 — — — 1Cleroidea Trogossitidae — — 1 1 — — 1 3 6Cucujoidea Erotylidae — — — 1 — — 2 6 9Cucujidae — — — — 1 — — 1 2Tenebrionoidea Mycetophagidae — — — — 1 — — 1 2Tenebrionidae — — — 3 — — — — 3Anthicidae — — 1 — — — — — 1Chrysomeloidea Cerambycidae — 1 9 8 3 — 8 8 37Curculionoidea Anthribidae — 4 — — 1 — — — 5Curculionidae 5 1 1 — — — 3 — 10Total 12 17 47 45 40 2 335 293 791CR, critically endangered; DD, data deficient; EN, endangered; EX, extinct; LC, least concern; LR/NT, lower risk–near threatened; NT, near threatened; VU, vulnerable. The order of families follows that in Table 11.1.11 Biodiversity ofColeoptera 3952005, Davis and Philips 2005, Bouchard et al. 2006, Talley and Holyoak 2006). Because the tax­onomy and distribution of most beetles remain unknown, we argue that the number of species currently listed as at‐risk represents a gross underestimation of the number that should be targeted for conservation.11.5 ConclusionsBeetles are a diverse group of arthropods that occur in most non‐marine habitats (and a few marine ones). Their influence on science and society is great. Beetles provide essential eco­logical services and are used as tools in many scientific endeavors, some with large effects on humans. On the other hand, beetles continue to have negative effects on vital industries such as agriculture and forestry. Studies on beetle bio­diversity and the conservation of their habitats are necessary to ensure the sustainability of nat­ural ecosystems and critical human activities.AcknowledgmentsWe thank P. 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