1 1 General Morphology, Classi�cation, and Biology of Cerambycidae Marcela L. Monné and Miguel A. Monné Universidade Federal do Rio de Janeiro Rio de Janeiro, Brasil Qiao Wang Massey University Palmerston North, New Zealand CONTENTS 1.1 Introduction ...................................................................................................................................... 2 1.2 De�nition and Morphology of the Family Cerambycidae ............................................................... 8 1.2.1 De�nition ............................................................................................................................. 8 1.2.2 General Morphology ........................................................................................................... 8 1.2.2.1 Adult ..................................................................................................................... 8 1.2.2.2 Immature Stages ................................................................................................ 28 1.3 Key to Subfamilies of the Family Cerambycidae .......................................................................... 29 1.3.1 Adults ................................................................................................................................. 36 1.3.2 Larvae ................................................................................................................................ 38 1.4 Diagnosis, Biodiversity, Distribution, and Biology of Subfamilies ............................................... 38 1.4.1 Subfamily Cerambycinae Latreille, 1802 .......................................................................... 40 1.4.1.1 Diagnosis ............................................................................................................ 40 1.4.1.2 Comments .......................................................................................................... 46 1.4.1.3 Diversity and Distribution .................................................................................. 46 1.4.1.4 Biology ............................................................................................................... 46 1.4.2 Subfamily Dorcasominae Lacordaire, 1868 ...................................................................... 46 1.4.2.1 Diagnosis ............................................................................................................ 46 1.4.2.2 Comments .......................................................................................................... 47 1.4.2.3 Diversity and Distribution .................................................................................. 47 1.4.2.4 Biology ............................................................................................................... 47 1.4.3 Subfamily Lamiinae Latreille, 1825.................................................................................. 47 1.4.3.1 Diagnosis ............................................................................................................ 47 1.4.3.2 Comments .......................................................................................................... 55 1.4.3.3 Diversity and Distribution .................................................................................. 56 1.4.3.4 Biology ............................................................................................................... 56 1.4.4 Subfamily Lepturinae Latreille, 1802 ............................................................................... 56 1.4.4.1 Diagnosis ............................................................................................................ 56 1.4.4.2 Comments .......................................................................................................... 58 1.4.4.3 Diversity and Distribution .................................................................................. 58 1.4.4.4 Biology ............................................................................................................... 58 2 Cerambycidae of the World 1.1 Introduction Cerambycidae Latreille, 1802, commonly known as longicorns, longhorns, longicorn beetles, longhorned beetles, longhorned borers, round-headed borers, timber beetles, or sawyer beetles, are among the most diverse and economically important families of Coleoptera. Taxonomic interest in the family has been fairly consistent for the past century, but the description of new taxa has accelerated in recent decades. The number of described cerambycid species in the world is about 36,300 in more than 5,300 genera (Tavakilian 2015). The adult body length ranges from less than 2 mm in Cyrtinus pygmaeus (Haldeman) (Linsley 1961) to greater than 170 mm in Titanus giganteus (L.) (Williams 2001). Cerambycids are widely distributed around the world—from sea level to 4,200 m above—wherever their host plants are found. Distribution and generic diversity of the world’s cerambycid subfamilies and tribes are shown in Table 1.1. The longicorn adults are free-living beetles that may or may not need to feed. They can live for a few days to a few months depending on whether they feed (Hanks 1999; Wang 2008). Cerambycids usually reproduce sexually but, in very rare cases—such as in some species of Kurarus Gressitt (Cerambycinae) (Goh 1977) and Cortodera Mulsant (Lepturinae) (Švácha and Lawrence 2014), they can reproduce parthenogenetically. Švácha and Lawrence (2014) suggested that at least in Cortodera, parthenogenesis probably is of recent origin because the female has a distinct spermatheca with a spermathecal gland. Mate location depends on the occurrence and status of larval hosts, adult food sources, and/or pheromones. Hanks (1999) predicted that the absence of feeding in the adult stage of many species is associated with the production of long-range pheromones, but the current knowledge shows that the use of volatile pheromones is widespread in ceramby- cids (see Chapter 5). The females lay their eggs on or near their hosts. The larvae of most cerambycid species feed on woody plants, but some select herbaceous hosts. The vast majority of species at the larval stage are living and feeding inside the plants although small minorities are free-living in soil and feed on plant roots. Many cerambycid larvae are dead plant feeders and play a major role in recycling dead plants; others attack living plants of different health states, ranging from stressed to healthy plants. To date, there are about 200 cerambycid species worldwide that have some economic impact on agriculture, forestry, and horticulture, causing billions of dollars of damage in production losses, environmental disasters, and management costs. They may damage plants by direct feeding and/or transmission of plant diseases. 1.4.5 Subfamily Necydalinae Latreille, 1825 ............................................................................. 59 1.4.5.1 Diagnosis ............................................................................................................ 59 1.4.5.2 Comments .......................................................................................................... 59 1.4.5.3 Diversity and Distribution .................................................................................. 59 1.4.5.4 Biology ............................................................................................................... 59 1.4.6 Subfamily Parandrinae Blanchard, 1845 ........................................................................... 60 1.4.6.1 Diagnosis ............................................................................................................ 60 1.4.6.2 Comments .......................................................................................................... 61 1.4.6.3 Diversity and Distribution .................................................................................. 61 1.4.6.4 Biology ............................................................................................................... 61 1.4.7 Subfamily Prioninae Latreille, 1802 ................................................................................. 61 1.4.7.1 Diagnosis ............................................................................................................ 61 1.4.7.2 Comments .......................................................................................................... 63 1.4.7.3 Diversity and Distribution .................................................................................. 63 1.4.7.4 Biology ............................................................................................................... 63 1.4.8 Subfamily Spondylidinae Audinet-Serville, 1832 ............................................................. 64 1.4.8.1 Diagnosis ............................................................................................................ 64 1.4.8.2 Comments .......................................................................................................... 65 1.4.8.3 Diversity and Distribution .................................................................................. 65 1.4.8.4 Biology ............................................................................................................... 65 Acknowledgments .................................................................................................................................... 66 References ................................................................................................................................................ 66 D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 3General Morphology, Classi�cation, and Biology of Cerambycidae TABLE 1.1 Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes Subfamilies and Tribes Biogeographic Regions No. Genera Cerambycinae Latreille, 1802 All biogeographic regions 1,757 Acangassuini Galileo & Martins, 2001 Neotropical 1 Achrysonini Lacordaire, 1868 All biogeographic regions 20 Agallissini Le Conte, 1873 Neotropical 3 Alanizini Di Iorio, 2003 Neotropical 1 Anaglyptini Lacordaire, 1868 All biogeographic regions 12 Aphanasiini Lacordaire, 1868 Afrotropical and Australian 6 Aphneopini Lacordaire, 1868 Australian 5 Auxesini Lepesme & Breuning, 1952 Afrotropical 8 Basipterini Fragoso, Monné & Campos Seabra, 1987 Neotropical 2 Bimiini Lacordaire, 1868 Australian and Neotropical 7 Bothriospilini Lane, 1950 Neotropical 11 Brachypteromatini Sama, 2008 Palaearctic 1 Callichromatini Swainson, 1840 All biogeographic regions 178 Callidiini Kirby, 1837 All biogeographic regions 38 Callidiopini Lacordaire, 1868 All biogeographic regions 62 Cerambycini Latreille, 1802 All biogeographic regions 99 Certallini Fairmaire, 1864 Palaearctic, Afrotropical, and Australian 9 Chlidonini Waterhouse, 1879 Afrotropical (Madagascar) 2 Cleomenini Lacordaire, 1868 Afrotropical and Oriental 23 Clytini Mulsant, 1839 All biogeographic regions 83 Compsocerini Thomson, 1864 All biogeographic regions 33 Coptommatini Lacordaire, 1869 Australian 1 Curiini LeConte, 1873 Neotropical 1 Deilini Fairmaire, 1864 Palaearctic and Australian 3 Dejanirini Lacordaire, 1868 Oriental 2 Diorini Lane, 1950 Neotropical 1 Distichocerini Pascoe, 1867 Australian 2 Dodecosini Aurivillius, 1912 Neotropical 4 Dryobiini Arnett, 1962 Nearctic and Neotropical 3 Eburiini Blanchard, 1845 Neotropical 23 Ectenessini Martins, 1998 Neotropical 12 Elaphidiini Thomson, 1864 Nearctic and Neotropical 91 Eligmodermini Lacordaire, 1868 Neotropical 5 Erlandiini Aurivillius, 1912 Neotropical 1 Eroschemini Lacordaire, 1868 Australian 2 Eumichthini Linsley, 1940 Nearctic 2 Gahaniini Quentin & Villiers, 1969 Afrotropical 1 Glaucytini Lacordaire, 1868 Oriental and Australian 18 Graciliini Mulsant, 1839 All biogeographic regions 22 Hesperophanini Mulsant, 1839 All biogeographic regions 85 Hesthesini Pascoe, 1867 Australian 1 Heteropsini Lacordaire, 1868 Neotropical and Australian 29 Hexoplini Martins, 2006 Neotropical 22 Holopleurini Chemsak & Linsley, 1974 Nearctic 1 Hyboderini Linsley, 1940 Nearctic and Neotropical 4 Hylotrupini Zagajkevich, 1991 Palaearctic 1 Ideratini Martins & Napp, 2009 Neotropical 1 (Continued) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 4 Cerambycidae of the World TABLE 1.1 (Continued) Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes Subfamilies and Tribes Biogeographic Regions No. Genera Lissonotini Swainson, 1840 Neotropical 1 Luscosmodicini Martins, 2003 Neotropical 1 Lygrini Sama, 2008 Afrotropical 1 Macronini Lacordaire, 1868 Australian 4 Megacoelini Quentin & Villiers, 1969 Afrotropical 2 Methiini Thomson, 1860 Oriental, Afrotropical, and Neotropical 19 Molorchini Gistel, 1848 All biogeographic regions 26 Mythodini Lacordaire, 1868 Oriental 4 Necydalopsini Lacordaire, 1868 Neotropical 12 Neocorini Martins, 2005 Neotropical 7 Neoibidionini Monné, 2012 Neotropical 55 Neostenini Lacordaire, 1868 Australian 4 Obriini Pascoe, 1871 All biogeographic regions 43 Ochyrini Pascoe, 1871 Australian 1 Oedenoderini Aurivillius, 1912 Afrotropical 1 Oemini Lacordaire, 1868 All biogeographic regions 101 Opsimini LeConte, 1873 Nearctic and Palaearctic 3 Oxycoleini Martins & Galileo, 2003 Neotropical 2 Paraholopterini Martins, 1997 Neotropical 1 Phalotini Lacordaire, 1868 Australian 4 Phlyctaenodini Lacordaire, 1868 Australian and Neotropical 17 Phoracanthini Newman, 1840 Australian 22 Phyllarthriini Lepesme & Breuning, 1956 Afrotropical 4 Piesarthriini McKeown, 1947 Australian 4 Piezocerini Lacordaire, 1868 Neotropical 19 Platyarthrini Bates, 1870 Neotropical 1 Plectogasterini Quentin & Villiers, 1969 Afrotropical 8 Plectromerini Nearns & Braham, 2008 Neotropical 1 Pleiarthrocerini Lane, 1950 Neotropical 1 Plesioclytini Wappes & Skelley, 2015 Nearctic 1 Proholopterini Monné, 2012 Neotropical 3 Protaxini Gahan, 1906 Oriental 1 Prothemini Lacordaire, 1868 Oriental 3 Psebiini Lacordaire, 1868 Afrotropical and Neotropical 24 Pseudocephalini Aurivillius, 1912 Australian and Neotropical 4 Pseudolepturini Thomson, 1861 Oriental 6 Psilomorphini Lacordaire, 1868 Australian 3 Pteroplatini Thomson, 1861 Afrotropical and Neotropical 10 Rhagiomorphini Newman, 1841 Australian 4 Rhinotragini Thomson, 1861 Neotropical 82 Rhopalophorini Blanchard, 1845 Nearctic, Neotropical, and Australian 29 Sestyrini Lacordaire, 1868 Oriental 2 Smodicini Lacordaire, 1868 Afrotropical, Nearctic, and Neotropical 8 Spintheriini Lacordaire, 1869 Australian 2 Stenhomalini Miroshnikov, 1989 Oriental 2 Stenoderini Pascoe, 1867 Australian and Oriental 10 Stenopterini Gistel, 1848 Palaearctic and Oriental 14 Strongylurini Lacordaire, 1868 Australian 6 (Continued) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 5General Morphology, Classi�cation, and Biology of Cerambycidae TABLE 1.1 (Continued) Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes Subfamilies and Tribes Biogeographic Regions No. Genera Tessarommatini Lacordaire, 1868 Australian 1 Thraniini Gahan, 1906 Oriental 3 Thyrsiini Marinoni & Napp, 1984 Neotropical 1 Tillomorphini Lacordaire, 1868 Nearctic, Neotropical, Oriental, and Australian 31 Torneutini Thomson, 1861 Neotropical 16 Trachyderini Dupont, 1836 All biogeographic regions 154 Tragocerini Pascoe, 1867 Australian 1 Trichomesiini Aurivillius, 1912 Australian 1 Trigonarthrini Villiers, 1984 Afrotropical 2 Tropocalymmatini Lacordaire, 1868 Australian 1 Typhocesini Lacordaire, 1868 Australian 4 Unxiini Napp, 2007 Neotropical 8 Uracanthini Blanchard, 1853 Australian 6 Vesperellini Sama, 2008 Palaearctic 1 Xystrocerini Blanchard, 1845 Afrotropical and Australian 2 Dorcasominae Lacordaire, 1868 Afrotropical, Oriental, and Palaearctic 95 Apatophyseini Lacordaire, 1869 Afrotropical, Oriental, and Palaearctic 90 Dorcasomini Lacordaire, 1868 Afrotropical, Oriental, and Palaearctic 5 Lamiinae Latreille, 1825 All biogeographic regions 2,964 Acanthocinini Blanchard, 1845 All biogeographic regions 386 Acanthoderini Thomson, 1860 All biogeographic regions 66 Acmocerini Thomson, 1864 Afrotropical 6 Acridocephalini Dillon & Dillon, 1959 Afrotropical 1 Acrocinini Swainson, 1840 Neotropical 1 Aderpasini Breuning & Teocchi, 1978 Afrotropical 1 Aerenicini Lacordaire, 1872 Neotropical 26 Agapanthiini Mulsant, 1839 All biogeographic regions 84 Amphoecini Breuning, 1951 Australian 2 Ancitini Aurivillius, 1917 Australian 1 Ancylonotini Lacordaire, 1869 Afrotropical, Oriental, and Palaearctic 36 Anisocerini Thomson, 1860 Neotropical 26 Apomecynini Thomson, 1860 All biogeographic regions 240 Astathini Thomson, 1864 Australian, Afrotropical, Oriental, and Palaearctic 23 Batocerini Thomson, 1864 Australian, Oriental, and Palaearctic 10 Calliini Thomson, 1864 Neotropical 40 Ceroplesini Thomson, 1860 Afrotropical, Oriental, and Palaearctic 88 Cloniocerini Lacordaire, 1872 Afrotropical 1 Colobotheini Thomson, 1860 Neotropical 12 Compsosomatini Thomson, 1867 Neotropical 13 Cyrtinini Thomson, 1864 Australian and Neotropical 16 Desmiphorini Thomson, 1860 All biogeographic regions 319 Dorcadionini Swainson, 1840 Palaearctic and Oriental 14 Dorcaschematini Thomson, 1860 Oriental and Australian 9 Elytracanthinini Bousquet, 2009 Neotropical 1 Enicodini Thomson, 1864 Australian and Oriental 27 Eupromerini Galileo & Martins, 1995 Neotropical 5 Forsteriini Tippmann, 1960 Neotropical 16 Gnomini Thomson, 1860 Australian, Oriental, and Palaearctic 4 (Continued) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 6 Cerambycidae of the World TABLE 1.1 (Continued) Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes Subfamilies and Tribes Biogeographic Regions No. Genera Gyaritini Breuning, 1950 Australian and Oriental 14 Heliolini Breuning, 1951 Australian 1 Hemilophini Thomson, 1868 Neotropical and Nearctic 127 Homonoeini Thomson, 1864 Australian, Oriental, and Palaearctic 22 Hyborhabdini Aurivillius, 1911 Oriental 1 Lamiini Latreille, 1825 Afrotropical, Australian, Oriental, and Palaearctic 48 Laticraniini Lane, 1959 Neotropical 2 Mauesiini Lane, 1956 Neotropical 4 Megabasini Thomson, 1860 Neotropical 1 Mesosini Mulsant, 1839 All biogeographic regions 99 Microcymaturini Breuning & Teocchi, 1985 Afrotropical 3 Moneilemini Thomson, 1864 Nearctic and Neotropical 1 Monochamini Gistel, 1848 All biogeographic regions 263 Morimonellini Lobanov, Danilevsky & Murzin, 1981 Palaearctic 1 Morimopsini Lacordaire, 1869 All regions except Nearctic 47 Nyctimeniini Gressitt, 1951 Australian and Oriental 1 Obereini Thomson, 1864 All regions except Neotropical 3 Oculariini Breuning, 1950 Afrotropical 2 Onciderini Thomson, 1860 Neotropical and Nearctic 81 Oncideropsidini Aurivillius, 1922 Oriental 1 Onocephalini Thomson, 1860 Neotropical 3 Onychogleneini Aurivillius, 1923 Oriental 1 Parmenini Mulsant, 1839 All biogeographic regions 87 Petrognathini Blanchard, 1845 Afrotropical and Oriental 10 Phacellini Lacordaire, 1872 Neotropical 7 Phantasini Kolbe, 1897 Afrotropical 3 Phrynetini Thomson, 1864 Afrotropical, Oriental, and Palaearctic 14 Phymasternini Teocchi, 1989 Afrotropical 1 Phytoeciini Mulsant, 1839 All biogeographic regions 32 Pogonocherini Mulsant, 1839 All biogeographic regions 33 Polyrhaphidini Thomson, 1860 Afrotropical and Neotropical 2 Pretiliini Martins & Galileo, 1990 Neotropical 1 Proctocerini Aurivillius, 1922 Afrotropical 1 Prosopocerini Thomson, 1864 Afrotropical 18 Pteropliini Thomson, 1860 All biogeographic regions 256 Saperdini Mulsant, 1839 All regions except Neotropical 154 Stenobiini Breuning, 1950 Afrotropical 7 Sternotomini Thomson, 1860 Afrotropical 20 Tapeinini Thomson, 1857 Neotropical and Oriental 2 Tetraopini Thomson, 1860 Nearctic and Neotropical 3 Tetraulaxini Breuning & Teocchi, 1977 Afrotropical 2 Tetropini Portevin, 1927 Palaearctic 2 Theocrini Lacordaire, 1872 Afrotropical 8 Tmesisternini Blanchard, 1853 Australian and Oriental 12 Tragocephalini Thomson, 1857 Afrotropical 63 Xenicotelini Matsushita, 1933 Oriental 1 Xenofreini Aurivillius, 1923 Neotropical 3 Xenoleini Lacordaire, 1872 Australian, Oriental, and Palaearctic 3 (Continued) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 7General Morphology, Classi�cation, and Biology of Cerambycidae TABLE 1.1 (Continued) Distribution and Generic Diversity of Cerambycid Subfamilies and Tribes Subfamilies and Tribes Biogeographic Regions No. Genera Xylorhizini Lacordaire, 1872 Afrotropical, Australian, Oriental, and Palaearctic 10 Zygocerini Thomson, 1864 Australian and Oriental 9 Lepturinae Latreille, 1802 All biogeographic regions 210 Desmocerini Blanchard, 1845 Nearctic 1 Encyclopini LeConte, 1873 Nearctic and Palaearctic 2 Lepturini Latreille, 1802 All biogeographic regions 140 Oxymirini Danilevsky, 1997 Palaearctic 1 Rhagiini Kirby, 1837 All biogeographic regions 53 Rhamnusiini Sama, 2009 Palaearctic and Oriental 2 Sachalinobiini Danilevsky, 2010 Nearctic and Palaearctic 1 Teledapini Pascoe, 1871 Oriental 3 Xylosteini Reitter, 1913 Palaearctic and Oriental 7 Necydalinae Latreille, 1825 Nearctic, Palaearctic, and Oriental 2 Necydalini Latreille, 1825 Nearctic, Palaearctic, and Oriental 2 Parandrinae Blanchard, 1845 All biogeographic regions 19 Erichsoniini Thomson, 1861 Neotropical 1 Parandrini Blanchard, 1845 All biogeographic regions 18 Prioninae Latreille, 1802 All biogeographic regions 302 Acanthophorini Thomson, 1864 Afrotropical 7 Aegosomatini Thomson, 1861 Afrotropical, Oriental, and Australian 20 Anacolini Thomson, 1857 Afrotropical, Oriental, and Neotropical 33 Cacoscelini Thomson, 1861 Afrotropical and Australian 5 Callipogonini Thomson, 1861 Afrotropical, Palaearctic, and Neotropical 17 Calocomini Galileo & Martins, 1993 Neotropical 1 Cantharocnemini Thomson, 1861 Afrotropical and Australian 6 Closterini, Lacordaire, 1868 Afrotropical, Australian, and Oriental 8 Ergatini Fairmaire, 1864 Afrotropical, Palaearctic, and Nearctic 5 Eurypodini Gahan, 1906 Palaearctic and Oriental 4 Hopliderini Thomson, 1864 Afrotropical 5 Macrodontiini Thomson, 1861 Neotropical 5 Macrotomini Thomson, 1861 All biogeographic regions 78 Mallaspini Thomson, 1861 Neotropical 10 Mallodonini Thomson, 1861 Afrotropical, Oriental, Nearctic, and Neotropical 10 Meroscelisini Thomson, 1861 Afrotropical, Australian, and Neotropical 21 Prionini Latreille, 1802 All biogeographic regions 50 Remphanini Lacordaire, 1868 Oriental 6 Solenopterini Lacordaire, 1868 Neotropical 7 Tereticini Lameere, 1913 Afrotropical and Australian 3 Vesperoctenini Vives, 2005 Neotropical 1 Spondylidinae Audinet-Serville, 1832 All biogeographic regions 32 Anisarthrini Mamaev & Danilevsky, 1973 Palaearctic 4 Asemini Thomson, 1861 All biogeographic regions 12 Atimiini LeConte, 1873 Nearctic, Neotropical, and Palaearctic 3 Saphanini Gistel 1848 Afrotropical and Nearctic 10 Spondylidini Audinet-Serville, 1832 Neotropical, Nearctic, and Palaearctic 3 D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 8 Cerambycidae of the World With the increase of international trade in recent years, many cerambycid species have been intercepted; some have become established outside their natural distribution range, causing serious problems globally (Haack et al. 2010; see Chapter 13). Linsley (1961, 1962a) and Wang (2008) summarize the general morphology and biology of the Cerambycidae. More recently, Švácha and Lawrence (2014) have made a very detailed treatment of the morphology and a general account of the ecology of the Cerambycidae. Ślipiński and Escalona (2013) gave a good introduction to the morphology and ecology of Australian cerambycids. In this chapter, we summa- rize the current knowledge about this family, including the de�nition and morphology, and a brief introduc- tion to the taxonomy, distribution, and general biology at the subfamily level. We aim to provide readers with a fundamental knowledge of cerambycids as well as a guide for those who may wish to consult speci�c chapters in this book where detailed treatments of Cerambycid biology and pest management are discussed. 1.2 Definition and Morphology of the Family Cerambycidae 1.2.1 Definition Traditionally, the family Cerambycidae had wider scope, including nine subfamilies: Anoplodermatinae, Aseminae, Cerambycinae, Lamiinae, Lepturinae, Parandrinae, Philinae, Prioninae, and Spondylidinae (Napp 1994). In the current classi�cation system (Bouchard et al. 2011; Monné 2012; Švácha and Lawrence 2014), Oxypeltinae, Vesperinae, and Disteniinae are considered independent families. We use the new system in this book and discuss eight subfamilies: Cerambycinae, Dorcasominae, Lamiinae, Lepturinae, Necydalinae, Parandrinae, Prioninae, and Spondylidinae. Table 1.1 summarizes the distri- bution and generic diversity of cerambycid subfamilies and tribes. 1.2.2 General Morphology The general morphology of Cerambycidae is extracted from Ślipiński and Escalona (2013) and Švácha and Lawrence (2014). 1.2.2.1 Adult 1.2.2.1.1 Diagnosis General external morphology of cerambycid adults is illustrated in Figures 1.1 and 1.2. Antennae usually �liform, elongate, and 11-segmented, rarely serrate and >12-segmented, usually inserted on pronounced tubercles; eyes usually emarginate; prothorax without pleural sutures; tibia with two distinct tibial spurs; tarsi usually pseudotetramerous with fourth tarsomere usually minute and concealed by third tarsomere; elytra usually covering abdomen; hind wings with a spur on radio-medial crossvein; abdomen usually with �ve visible sternites, �fth sternite entire. 1.2.2.1.2 Description 1.2.2.1.2.1 Head The head is prognathous and more or less horizontal in the Parandrinae (Figures 1.3 and 1.4). It is produced anteriorly to form a short to moderately long muzzle in some Lepturinae (Figures 1.5 and 1.6), Dorcasominae, and Cerambycinae, inclined anteriorly in the Spondylidinae, and is vertical or retracted, with the genal line directed posteriorly, in the Lamiinae (Figure 1.7). The eyes are entire in the Parandrinae (Figure 1.3), most Lepturinae, and some Prioninae; feebly emarginate in the Spondylidinae (Figure 1.8) and most Prioninae (Figure 1.9); emarginate to entire in the Dorcasominae; and usually are deeply emarginate and reniform in the Cerambycinae (Figure 1.10) and Lamiinae (Figure 1.7); although occasionally they are divided—as in Tetraopes Schönherr—or lacking the upper lobe—as in Tillomorpha Blanchard. The facets of the eyes are large and coarse in the Parandrinae, most Prioninae, and some Asemini and Cerambycinae; usually, they are �ner in the Lepturinae, Lamiinae, and more specialized Cerambycinae. The antennae usually have 11 antennomeres (Figures 1.1 and 1.2) that are inserted near the base of the mandibles in the Parandrinae (Figure 1.3), Prioninae (Figure 1.9), and in some Spondylidinae; D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 9General Morphology, Classi�cation, and Biology of Cerambycidae Head Scape Pedicel Claw Protarsomere V Mesofemur Metafemur Metatibia MetatarsomeresElytral apex Elytral suture Elytrum Scutellum Pronotum Eye FIGURE 1.1 General morphology, dorsal view of Trachyderes succinctus (L.) (Cerambycinae). Labium Mandible Prosternum Procoxal cavity Mesepisternum Mesepimerum Metepisternum Metacoxal cavity Sternite Metacoxa Metasternum Mesosternal process Prosternal process FIGURE 1.2 General morphology, ventral view of Trachyderes succinctus (L.) (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 10 Cerambycidae of the World 1m m FIGURE 1.4 Head, dorsal view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.5 Head, dorsal view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1 m m FIGURE 1.3 Head, lateral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-039.jpg&w=179&h=170 11General Morphology, Classi�cation, and Biology of Cerambycidae 1m m FIGURE 1.6 Head, lateral view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.7 Head, lateral view of Estola obscura Thomson (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.8 Head, lateral view of Asemum striatum (L.) (Spondylidinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-058.jpg&w=210&h=136 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-059.jpg&w=142&h=180 12 Cerambycidae of the World are near the eyes in the Asemini and Lepturinae (Figure 1.6); and are usually more or less embraced by the eyes in the Cerambycinae (Figure 1.10) and Lamiinae (Figure 1.7). In some diurnal Lamiinae (e.g., Octotapnia Galileo & Martins and Pseudotapnia Chemsak & Linsley) and Prioninae, the antennae may have fewer antennomeres. In some Lamiinae (e.g., Paratenthras Monné), the �rst three antennomeres are long, whereas the remaining �agella are reduced and sometimes moniliform. The number of antennomeres may be 12 in a number of unrelated groups and more than 12 in a few Cerambycinae and Prioninae (up to 30 in some species of Prionus Müller). The antennal structure is similar between sexes in the Parandrinae, Spondylidinae, and Lepturinae, and strikingly dissimilar in many Prioninae and in most Cerambycinae and Lamiinae. In the Parandrinae and Spondylidinae, differentiation of antennomeres is not well marked; the scape is short, the second antennomere is not greatly reduced in size, half as long as, or subequal to the third antennomere, and the segments that follow are subequal in length. In the remaining subfamilies, the scape is usually more elongate, the  second segment is greatly reduced, and the following antennomeres are unequal in length— with the third usually greatly elongated and those that follow diminishing to the ultimate antenno- mere. The antennal segments are glabrous in the Parandrinae, Prioninae, and Spondylidinae, and are pubescent in other subfamilies. The labrum is fused with the epistoma in the Parandrinae and Prioninae but free in other subfami- lies. The mandibles are acute in all of the Cerambycidae; large and often toothed in the Parandrinae (Figure 1.4) and Prioninae (Figure 1.11); long, slender, and untoothed in the Spondylidinae; shorter in most other groups; and are provided with a dense fringe of hairs in the inner margin of the Dorcasominae and Lepturinae. The maxillae are typically bilobed; the inner lobe is obsolete in the Parandrinae (Figure 1.12) and Prioninae (Figure 1.13). The ultimate segment of the palpi (both 1m m FIGURE 1.9 Head, lateral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.10 Head, lateral view of Achryson surinamum (L.) (Cerambycinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 13General Morphology, Classi�cation, and Biology of Cerambycidae maxillary and labial) is pointed at the apex in the Lamiinae (Figures 1.14 and 1.15) and truncated (Figures 1.16 through 1.19) in other subfamilies. The submentum projects between the bases of the maxillae in the Lepturinae; is short in many Cerambycinae; and is absent in the Parandrinae, Prioninae, and Spondylidinae. The mentum is distinctly transverse in the Parandrinae (Figure 1.20), Prioninae, and Spondylidinae, and trapezoidal in the Lepturinae, Cerambycinae (Figure 1.18), and Lamiinae (Figure 1.14). The ligula is corneous in the Parandrinae and Spondylidinae, and membra- nous or coriaceous in the Lepturinae, Cerambycinae (Figure 1.18) (except Oemini and Methini), and Lamiinae (Figure 1.14). 1m m FIGURE 1.11 Head, dorsal view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.12 Maxilla, ventral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permis- sion from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-153.jpg&w=105&h=238 14 Cerambycidae of the World 1.2.2.1.2.2 Thorax The prothorax bears lateral carinae in the Parandrinae (Figure 1.21) and Prioninae (Figures 1.22 and 1.23), which are lacking in other subfamilies (Figures 1.24 through 1.26). The pro- coxae are strongly transverse in the Parandrinae and Prioninae, less so in some Spondylidinae—such as Asemini, subconical in the rest Spondylidinae, conical in the Lepturinae, and usually rounded in the Cerambycinae  and  Lamiinae. The procoxal cavities are closed behind in some Parandrinae, in some Spondylidinae, and in most Lamiinae (Figure 1.26); wide open in the Prioninae (Figure 1.23), Asemini, and most Lepturinae (Figure 1.25); and open or closed in the Cerambycinae. The scutellum is visible, some- times well developed (Figure 1.1) and usually is not abruptly elevated, anteriorly �at, or separated from 1m m FIGURE 1.13 Maxilla, ventral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.14 Labium, ventral view of Estola obscura Thomson (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-154.jpg&w=100&h=234 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-155.jpg&w=168&h=182 15General Morphology, Classi�cation, and Biology of Cerambycidae 1m m FIGURE 1.15 Maxilla, ventral view of Estola obscura Thomson (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.16 Maxilla, ventral view of Asemum striatum (L.) (Spondylidinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.17 Maxilla, ventral view of Necydalis major L. (Necydalinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-156.jpg&w=131&h=163 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-157.jpg&w=153&h=128 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-158.jpg&w=136&h=193 16 Cerambycidae of the World 1m m FIGURE 1.18 Labium, ventral view of Rhopalophora collaris (Germar) (Cerambycinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.19 Maxilla, ventral view of Trachyderes succinctus (L.) (Cerambycinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.20 Labium, ventral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permis- sion from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-159.jpg&w=146&h=168 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-160.jpg&w=124&h=203 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-161.jpg&w=223&h=102 17General Morphology, Classi�cation, and Biology of Cerambycidae 1m m FIGURE 1.21 Prothorax, lateral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permis- sion from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.22 Prothorax, lateral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.23 Prosternum, ventral view of Mallodon spinibarbis (L.) (Prioninae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-162.jpg&w=156&h=167 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-163.jpg&w=178&h=173 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-164.jpg&w=202&h=131 18 Cerambycidae of the World 1m m FIGURE 1.24 Prothorax, lateral view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.25 Prosternum, ventral view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.26 Prosternum, ventral view of Adesmus hemispilus Germar (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-165.jpg&w=177&h=155 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-166.jpg&w=177&h=163 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-167.jpg&w=168&h=153 19General Morphology, Classi�cation, and Biology of Cerambycidae mesoscutum by an impression. The mesonotum lacks a stridulatory plate in the Parandrinae (Figure 1.27), Prioninae, and some Spondylidinae; has a divided stridulatory plate in the Dorcasominae (if present), Asemini (Spondylidinae) (Figure 1.28), and Lepturinae (Figure 1.29), and an undivided stridulatory plate in the Necydalinae (Figure 1.30), Cerambycinae (Figure 1.31), and Lamiinae (Figure 1.32). Legs mostly are cursorial (Figure 1.1) and usually moderately long in most longicorn beetles but can be very long in some species such as males of lamiine Gerania (Audinet-Serville); fore legs are enlarged in some (particularly males) Prioninae and Lamiinae and extremely long in the lamiine Acrocinus Illiger (fore femora in large males can be as long as body), where they reportedly are used for traversing tree branches; hind legs may be enlarged, such as metafemora in the male cerambycine Utopia Thomson, or plate-like tibial extensions in some Cerambycinae, but are never adapted for jumping. The tibia usually has two spurs at the terminal end (Figure 1.33). The legs exhibit an oblique groove along the inner side of the protibiae and a notch or groove on the outer face of the mesotibiae in the Lamiinae; these grooves and notches are lacking in other subfamilies. The tarsi are distinctly pentamerous without pubescent ventral 1m m FIGURE 1.27 Mesonotum, dorsal view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.28 Mesonotum, dorsal view of Asemum striatum (L.) (Spondylidinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-168.jpg&w=131&h=201 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-169.jpg&w=198&h=153 20 Cerambycidae of the World 1m m FIGURE 1.29 Mesonotum, dorsal view of Leptura rubra L. (Lepturinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) FIGURE 1.30 Mesonotum, dorsal view of Necydalis major L. (Necydalinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.31 Mesonotum, dorsal view of Rhopalophora collaris (Germar) (Cerambycinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-170.jpg&w=155&h=173 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-171.jpg&w=119&h=170 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-172.jpg&w=141&h=140 21General Morphology, Classi�cation, and Biology of Cerambycidae pads in the Parandrinae (Figure 1.34), although they are pseudotetramerous with ventral pads in the Prioninae, Lepturinae, Cerambycinae, Spondylidinae (Figure 1.35), and Lamiinae. The third tarsomere is simple in the Parandrinae but dilated in the remaining subfamilies (Figure 1.35). The tarsal claws are appendiculate or cleft in the most specialized Lamiinae but simple in all other subfamilies. The hind wing usually has a moderately to very long apical �eld (though this is short in some very large forms, such as Titanus Audinet-Serville) with two more or less complete radial extensions converg- ing and then diverging to form a scissor-like �gure, with a dark sclerite apicad of radial cell and a subtri- angular sclerite crossing r4. The radial cell often is well developed and more or less elongate (although sometimes it is short and broad or lacking basal limit). Cross-vein r3 is slightly to strongly oblique and sometimes is absent. The basal portion of radius posterior (RP) is long to very short and not surpassing r4. The medial �eld usually has four or �ve free veins (sometimes with three or, rarely, fewer) and always lacks medial �eck. The wedge cell is well developed in almost all Prioninae and some Lepturinae and Spondylidinae and is absent in all other subfamilies. If the elytra are shortened in macropterous forms, the hind wings are exposed (often giving the beetles a hymenopteran appearance) and their apex is then sometimes not folded (all Necydalinae, Figures 1.36 and 1.37). The hind wings are highly reduced or disappear in numerous Cerambycinae (such as males of Torneutes Reich), Lamiinae (usually both sexes) and Prioninae (more often only females), Lepturinae, and Spondylidinae (both sexes of Michthisoma LeConte); in some taxa, the beetles apparently are �ightless even if wings are present. 1.2.2.1.2.3 Abdomen The abdomen usually has �ve free, visible ventrites (belonging to segments III–VII; sternites 1 and 2 form the posterointernal wall of metacoxal acetabula) (Figure 1.2), with the �rst usually not much longer than the second; rarely, it is almost as long as the remaining combined (females of the ceram- bycine Obriini). The intercoxal process is acute to broadly rounded or angulate—or absent, with the medial part of reduced sternum II visible between the hind coxae (Necydalinae and some slender wasp-mimicking 1m m FIGURE 1.32 Mesonotum, dorsal view of Adesmus hemispilus Germar (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.33 Protibia, lateral view of Spondylis buprestoides (L.) (Spondylidinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-173.jpg&w=161&h=149 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-174.jpg&w=288&h=77 22 Cerambycidae of the World Cerambycinae and the telescoped, with segment II forming a petiolus-like basal piece). The abdominal tergites 1–6 are semisclerotized. Functional spiracles are present on each side of abdominal segments I–VII (the �rst pair is very large, particularly in �ying forms), and spiracles VIII are rudimentary and closed. 1.2.2.1.2.4 External Morphology of Terminalia Male terminalia (Figures 1.38 through 1.40) consist of three abdominal segments. The anterior edge of sternite VIII (Figure 1.38) usually bears a median strut (that is rudimentary or absent in some taxa); the anterior edge of sternite IX has spiculum gastrale; tergites IX and X are fused together and usually membranous. The anterior edge of tegmen (Figure 1.39) 1m m FIGURE 1.34 Metatarsus, lateral view of Parandra (Parandra) glabra (De Geer) (Parandrinae). (Reprinted with permis- sion from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) 1m m FIGURE 1.35 Metatarsus, dorsal view of Spondylis buprestoides (L.) (Spondylidinae). (Reprinted with permission from C. J. B. Carvalho, editor. Napp, D. S., Rev. Bras. Entomol., 38, 265–419, 1994.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-175.jpg&w=60&h=209 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-176.jpg&w=96&h=209 23General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.36 Necydalis major L. (Necydalinae). FIGURE 1.37 Ulochaetes leoninus LeConte (Necydalinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-177.jpg&w=86&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-178.jpg&w=184&h=269 24 Cerambycidae of the World has a single or no strut; the parameres usually are fused to phallobase and free from one another, but they are more or less completely fused in some Cerambycinae (such as the Molorchini–Obriini complex— very short in some and nearly absent in the Neotropical Ectenessini). The aedeagus is cucujiform and symmetrical, but usually is rotated to one side in the abdominal cavity when at rest. Surrounding struc- tures therefore may not be entirely symmetrical. The anterior edge of the aedeagus (Figure 1.40) almost 1m m FIGURE 1.38 Male sternite VIII of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) 1m m FIGURE 1.39 Tegmen of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-179.jpg&w=141&h=225 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-180.jpg&w=108&h=229 25General Morphology, Classi�cation, and Biology of Cerambycidae always has paired struts; the internal sac (endophallus) (Figure 1.40) may bear distinctive sclerotized structures such as asperities, paired or unpaired sclerites, or longitudinal sclerotized rods. The terminalia of female cerambycids (Figures 1.41 and 1.42) follow the same structural plan as the male, with sclerites sequentially distributed along a more or less membranous tube, which is kept invagi- nated at rest. Segment VIII is entirely contained within the partially double-walled segment VII (where the �rst invagination occurs). Sternite VIII (Figure 1.41) bears a long ventral apodeme that is closely related with the ovipositor length among different taxa. Sternite and tergite VIII are mostly partly des- clerotized and tend to fuse into a tube enclosing the “anus–ovipositor” complex, sometimes protruding from the abdomen, either “naked” or protected by posterior sternal and tergal projections of segment VII (e.g., some Acanthocinini of Lamiinae). The ovipositor (Figure 1.42) usually is long but secondarily shortened and modi�ed—particularly in some Cerambycinae (such as Trachyderini) ovipositing on the host surface; the paraprocts are short without baculi (in all Lamiinae) and �exible with subapical styli; in some groups, the apex of the ovipositor is sclerotized with styli being lateral or laterodorsal and often reduced or virtually inbuilt in coxitis (this type occurs in several subfamilies depending on biology but is common in the Prioninae and universal in the Parandrinae). One or two pairs of glandular integumental invaginations often are present at the base of the ovipositor on both sides of anus. 1.2.2.1.2.5 Reproductive System Testes (Figure 1.43) consist of one to several pairs of testicular lobes, with each lobe having a number of radially arranged testicular follicles. The basal parts of vasa deferen- tia may be broadened into seminal vesicles. Usually, there are two pairs (or at least one pair) of accessory glands at or before the fusion of vasa deferentia. Ducts are more or less completely paired (mostly up to paired gonopores on the internal sac) in Lamiini and several related tribes of Lamiinae. Primary gono- pore seldom projects into a long sclerotized �agellum. Ovaries (Figure 1.44) are paired, each with a variable number (up to several tens) of ovarioles. There is a single more or less sclerotized spermatheca of simple shape (often an elongate, curved capsule bridged 1m m FIGURE 1.40 Aedeagus and internal sac of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-181.jpg&w=120&h=264 26 Cerambycidae of the World 1m m FIGURE 1.41 Female terminalia of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) 1m m FIGURE 1.42 Ovipositor of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-183.jpg&w=155&h=259 27General Morphology, Classi�cation, and Biology of Cerambycidae 1m m FIGURE 1.43 Male reproductive system of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) 1m m FIGURE 1.44 Female reproductive system of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-184.jpg&w=226&h=265 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-185.jpg&w=181&h=260 28 Cerambycidae of the World by spermathecal compressor) with a distinct, sometimes very large, spermathecal gland attached to it. A bursa copulatrix usually is present in the form of a soft diverticulum of various sizes and locations; the spermathecal duct arises on its base. 1.2.2.1.2.6 Digestive System The adult gut may be primitively rather reduced. In prionines and some cerambycines, the midgut is short and reduced with the posterior midgut being extremely reduced and thread-like (slightly less so in some �oricolous Clytini of the Cerambycinae), suggesting adult aphagy. However, the gut is long and well developed in the Lamiinae whose adults feed extensively. The digestive tube (Figure 1.45) usually does not have a distinct crop/proventriculus. The anterior midgut in some taxa (Necydalinae, Spondylidinae, and most Lepturinae) produces morphologically dis- tinct mycetomes in the form of gut wall diverticula whose cells harbor intracellular yeast-like symbionts. The posterior midgut often bears numerous small, scattered evaginated crypts. Six cryptonephridial Malpighian tubules enter the gut separately in two clusters of three. 1.2.2.2 Immature Stages The following morphological descriptions are based on Butovitsch (1939), Duffy (1953, 1957, 1960, 1968), Gardiner (1966), and Švácha and Lawrence (2014). 1.2.2.2.1 Eggs The eggs (Figure 1.46) are elongate, ovoid, or fusiform and often have thin, �exible chorion so that their shape can adapt to the tight spaces in which they usually are laid. A female can lay a dozen to several hundred eggs in her lifetime. They usually hatch in a few days to a few weeks after oviposition, depend- ing on species and climates. In some lamiine species, the larvae may overwinter within the chorion, particularly if the eggs are laid late in the season. 1.2.2.2.2 Larvae The larvae are soft-bodied, eucephalic, oligopodous to apodous, prognathous, more or less elongate, and subcylindrical to dorsoventrally depressed (Figures 1.47 through 1.58). Their body shape and mechanics largely depend on hemolymph pressure. The cranium (particularly its anterior part, which supports mouthparts, often called the “mouth- frame”) may be strongly sclerotized and pigmented, whereas the body generally is soft and white to yellow (Figures 1.51 through 1.54). In rare cases, the body can be grayish with some prothoracic regions and the abdominal end sclerotized and pigmented. The skin of the prothorax is not attached to the submentum. The ventral mouthparts are protracted; the mandibles (Figure 1.59) lack a molar tooth or other appendage; the labium bears a setose ligula, and gula and hypostoma are present. The abdomen, at least dorsally, has more or less retractile and often characteristically sculptured, protu- berances called ambulatory ampullae, providing support in galleries; abdominal segments 1 to 6 or 7 have dorsal ampullae. The spiracular system is peripneustic, with one pair of functional spiracles on the mesothorax (Figure 1.60) and one pair on each of eight abdominal segments. The digestive system is similar to that of adults. 1.2.2.2.3 Pupae The pupae (Figure 1.61a and b) are similar to adults in size, shape, and proportions of cephalic and thoracic appendages. Secondary sexual differences in adults generally are evident in the pupae. They are adecticous, exarate, and generally soft and pale (except for some special structures like spines or gin  traps), with a strongly ventrally bent head so that mouthparts point caudally (except for some Prioninae). The body usually is waxy or milky white to testaceous, often with scattered setae or spinose areas or combinations of both. The antennae extend at least as far as the mesothorax but generally to the abdominal segments, where they are nearly always curved downward beneath the body. The elytra are always glabrous (except Acanthocinini). The abdomen usually has nine movable segments, with the tenth (and occasionally D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 29General Morphology, Classi�cation, and Biology of Cerambycidae the ninth) being telescoped within the preceding segments. Abdominal segments 7 and 8 usually are more elongate than the preceding ones but sometimes considerably produced. The abdomen has �ve to seven pairs of functional spiracles. Segment 9 often ends in a vertical or horizontal spine or process or with a pair of incurved or outwardly curved urogomphi. Some prionines (tribes Callipogonini and Macrotomini) have paired paramedian gin-traps. The legs often have subapical setae on the femora and sometimes one or two setae on the tarsi. 1.3 Key to Subfamilies of the Family Cerambycidae The key to adults is based on the work of Linsley (1962b), Ślipiński and Escalona (2013), and Švácha and Lawrence (2014). The key to larvae is based on Duffy (1953, 1957, 1960, 1968), Švácha and Danilevsky (1987, 1988, 1989), and Švácha and Lawrence (2014). 1m m FIGURE 1.45 Digestive system of Oxymerus luteus luteus (Voet) (Cerambycinae). (Reprinted with permission from W. F. de Azevedo, Jr., editor. Moura, L. A., and A. F. Franceschini, Biociências, 2, 135–143, 1994.) 1m m FIGURE 1.46 Egg of Hedypathes betulinus (Klug) (Lamiinae). (Reprinted with permission from C. J. B. Carvalho, editor. Galileo, M. H. M., et al., Rev. Bras. Entomol., 37, 705–715, 1993.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-186.jpg&w=66&h=260 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-187.jpg&w=179&h=84 30 Cerambycidae of the World 1m m (a) (b) FIGURE 1.47 Larvae of Acanthoderes (Psapharochrus) melanosticta White (Lamiinae), dorsal view (a) and lateral view (b). (Reprinted with permission from C. J. B. Carvalho editor. Mermudes, J. R. M., and M. L. Monné, Rev. Bras. Entomol., 45, 331–334, 2001.) 5 m m FIGURE 1.48 Larva of Parandra sp. (Parandrinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-188.jpg&w=96&h=226 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-189.jpg&w=60&h=225 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-190.jpg&w=60&h=264 31General Morphology, Classi�cation, and Biology of Cerambycidae 5m m FIGURE 1.49 Larva of Parandra sp. (Parandrinae), lateral view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) 0, 3 m m FIGURE 1.50 Larva of Acyphoderes aurulenta (Kirby) (Cerambycinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-191.jpg&w=67&h=260 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-192.jpg&w=74&h=260 32 Cerambycidae of the World FIGURE 1.51 Larva of Tsivoka simplicicollis (Gahan) (Dorcasominae), laterodorsal view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) FIGURE 1.52 Larva of Tsivoka simplicicollis (Gahan) (Dorcasominae), ventral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-193.jpg&w=68&h=229 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-194.jpg&w=62&h=229 33General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.53 Larva of Judolia sexmaculata (L.) (Lepturinae), ventral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) FIGURE 1.54 Larva of Dinoptera collaris (L.) (Lepturinae), dorsal view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-195.jpg&w=105&h=229 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-196.jpg&w=104&h=229 34 Cerambycidae of the World FIGURE 1.56 Larva of Prionus coriarius (L.) (Prioninae), ventral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) FIGURE 1.55 Larva of Prionus coriarius (L.) (Prioninae), lateral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-197.jpg&w=63&h=229 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-198.jpg&w=56&h=229 35General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.57 Larva of Atimia okayamensis Hayashi (Spondylidinae), lateral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) FIGURE 1.58 Larva of Arhopalus rusticus (L.) (Spondylidinae), ventral view. (Reprinted with permission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3  Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-199.jpg&w=62&h=229 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-200.jpg&w=65&h=229 36 Cerambycidae of the World 1.3.1 Adults 1. Tarsi distinctly pentamerous (Figure 1.34); lateral pronotal carinae entire and simple (Figure 1.21) .......................................................................................................... Parandrinae a. Tarsi pseudotetramerous (Figure 1.35); lateral pronotal carinae absent (Figure 1.24), or present, often dentate or spinose (Figures 1.22 and 1.23) .................................................. 2 0. 5m m FIGURE 1.60 Thoracic spiracle of the larva of Acyphoderes aurulenta (Kirby) (Cerambycinae). (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) 1m m FIGURE 1.59 Mandible of the larva of Acyphoderes aurulenta (Kirby) (Cerambycinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-201.jpg&w=124&h=200 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-202.jpg&w=184&h=200 37General Morphology, Classi�cation, and Biology of Cerambycidae 2. Head vertical or retracted (Figure 1.7), genal margin always directed posteriorly; protibiae with mesial sinus; mesotibiae usually notched or grooved externally; last segment of maxillary palpi usually pointed at apex (Figures 1.14 and 1.15) ......................................................... Lamiinae a. Head usually prognathous or weakly de�exed (Figure 1.10), genal margin never directed posteriorly; protibiae without mesial sinus; mesotibiae never notched or grooved externally; last segment of maxillary palpi obtuse or truncate at apex (Figures 1.13 and 1.16 through 1.19) ................................................................................................................................... 3 3. Pronotum with elevated, often dentate or spinose lateral carinae (Figures 1.22 and 1.23); labrum fused with epistoma; inner lobe of maxillae obsolete (Figure 1.13); procoxae strongly transverse (Figure 1.23); mesoscutum without a striated stridulatory area .............. Prioninae a. Pronotum without distinct lateral carinae; labrum free; inner lobe of maxillae usually well developed (Figures 1.16, 1.17, and 1.19); procoxae rarely transverse, usually rounded; mesoscutum with a �nely striated stridulatory area (Figures 1.28 through 1.30) or without a stridulatory area ............................................................................................................... 4 4. Mesoscutum without a median endocarina (Figure 1.30); elytra shortened, covering only the pterothorax, exposed hind wings with unfolded apex (Figures 1.36 and 1.37) ..... Necydalinae a. Mesoscutum with a median endocarina (Figures 1.28 and 1.29); elytra usually well devel- oped, hind wings folded at apex ........................................................................................ 5 5. Head with region behind eyes usually having prominent temples followed by a constricted neck (Figures 1.5 and 1.6); procoxae conical, prominent, and strongly projecting below prosternal process ..................................................................................................................... Lepturinae a. Head may be gradually narrowing to abruptly constricted behind eyes, without prominent temples, followed by a constricted neck; procoxae of variable shape usually subglobular, seldom strongly projecting below prosternal process ........................................................ 6 1m m (a) 1m m (b) FIGURE 1.61 Pupae of Acanthoderes (Psapharochrus) melanosticta (White) (Lamiinae), dorsal view (a) and ventral view (b). (Reprinted with permission from C. J. B. Carvalho, editor. Mermudes, J. R. M., and M. L. Monné, Rev. Bras. Entomol., 45, 331–334, 2001.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 38 Cerambycidae of the World 6. Mandibula with incisor edge without fringe of hairs; hind wing with edge cell ................................................................................................................... Spondylidinae a. Mandibula with incisor edge usually with fringe of long hairs; hind wing without edge cell ............................................................................................................................................. 7 7. Mesonotum with an undivided stridulatory plate (Figure 1.31); notosternal suture rarely com- plete, usually indistinct or incomplete anteriorly or absent; empodium usually small or indis- tinct .................................................................................................................... Cerambycinae a. Mesonotum with a divided stridulatory plate; notosternal suture may be relatively distinct and complete; empodium indistinct ............................................................. Dorcasominae 1.3.2 Larvae 1. Clypeus (Figure 1.62) very narrow, with only slender basal arms reaching to mandibular articulations; these arms may be more or less sclerotized and fused with epistomal margin. Mandibular apex and dorsal angle lacking; mandibles (Figure 1.59) short and apically rounded, spoon-like .................................................................................... Cerambycinae (Figure 1.50) a. Clypeus more or less trapezoidal, �lling entire space between dorsal mandibular articula- tions. Mandibles with distinct apex and more or less distinct dorsal angle ...................... 2 2. Legs absent, or present with only two minute segments visible under high magni�cation. Cardo extremely reduced, labiomaxillary base �rmly attached to cranium along whole width; maxil- lae movable only from stipes ..................................................... Lamiinae (Figure 1.47a and b) a. Distinct four-segmented legs (Figure 1.65) present though may be strongly reduced and inconspicuous. Free movable cardo present ....................................................................... 3 3. Main antennal sensillum �at, rarely convex, never conical ..................................................... 4 a. Main antennal sensillum prominent and conical ............................................................... 5 4. Basal half of pronotum more or less roughly asperate. Labrum cordate (Figures 1.63 and 1.64), very long. Epistomal, frontal, and postcondylar carinae absent ..... Parandrinae (Figures 1.48 and 1.49) a. Body without coarse asperities. Labrum never as long as in Parandra. Distinct epistomal, frontal, and postcondylar carinae often present ........... Prioninae (Figures 1.55 and 1.56) 5. Pretarsus without setae. Abdominal epipleurum protuberant on segments 7–9. Lateral furrows of pronotum long and distinct ................................................................................................... 6 a. Pretarsus with distinct setae. Abdominal epipleurum protuberant on at least segments 6–9. Lateral furrows of pronotum rarely long and distinct ......................................... 7 6. Large postnotal fold present. Urogomphi absent. Dorsal ampullae with one lateral impression on each side ................................................................... Dorcasominae (Figures 1.51 and 1.52) a. Postnotum absent. Urogomphi present or absent. Dorsal ampullae with two lateral impres- sions on each side ................................................... Spondylidinae (Figures 1.57 and 1.58) 7. Dorsal ampullae with two broadly separate lateral impressions on each side (Figure 1.66) Prothoracic lateropresternum largely microspiculate ............................................ Necydalinae a. Dorsal ampullae with one lateral impression. Prothoracic lateropresternum microspiculate at most along anterior margin ..................................... Lepturinae (Figures 1.53 and 1.54) 1.4 Diagnosis, Biodiversity, Distribution, and Biology of Subfamilies Phylogenetic relationships within the Cerambycidae are still highly controversial (e.g., Wang and Chiang 1991; Napp 1994; Švácha and Lawrence 2014). As a result, we order subfamilies alphabetically in this section. Morphological features of subfamilies mainly are extracted from Linsley (1962a, 1962b, 1963, 1964), Linsley and Chemsak (1972, 1984, 1995), Chemsak (1996), Ślipiński and Escalona (2013), and Švácha and Lawrence (2014). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 39General Morphology, Classi�cation, and Biology of Cerambycidae 1m m FIGURE 1.62 Head of the larva of Acyphoderes aurulenta (Kirby) (Cerambycinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) 2. 5 m m FIGURE 1.63 Head of the larva of Parandra sp. (Parandrinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) 0. 5 m m FIGURE 1.64 Labrum of the larva of Parandra sp. (Parandrinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-207.jpg&w=174&h=150 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-208.jpg&w=159&h=161 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-209.jpg&w=142&h=150 40 Cerambycidae of the World 1.4.1 Subfamily Cerambycinae Latreille, 1802 1.4.1.1 Diagnosis Small to large, elongate beetles (Figures 1.67 through 1.76). Head (Figure 1.10) subvertical, scarcely narrowed behind eyes; antennae inserted high on frons between eyes, usually very elongate, second antennomere short; eyes reniform, usually embracing antennal insertion; mandibles acute without molar plate; incisor edge with or without pubescent fringe; maxillae and labium variable, lacinia usu- ally well developed (Figure 1.19); mentum usually trapezoidal (Figure 1.18); submentum sometimes produced between bases of maxillae as a short process. Pronotum without lateral margin; procoxae rarely prominent, usually rounded, cavities variable; notosternal suture rarely complete, usually indistinct or incomplete anteriorly, or absent; mesoscutum with a median endocarina; stridulatory plate, when present, undivided (Figure 1.31). Elytra usually not abbreviated; hind wings without closed cell in anal sector, radial cell closed. Legs moderately long; protibiae without mesial sinus; tarsi pseudotetramerous, padded beneath, third tarsomere dilated, bilobed concealing minute fourth tarsomere, empodium small or absent. 1m m FIGURE 1.65 Proleg of the larva of Parandra sp. (Parandrinae), dorsal view. (Reprinted from Costa, C., et al., Larvas de Coleoptera do Brasil, Museu de Zoologia, Universidade de São Paulo, São Paulo, 1988. With permission.) FIGURE 1.66 Larva of Necydalis major L. (Necydalinae), detail of ventral ambulatory ampulla 4. (Reprinted with per- mission from Petr Švácha, owner. Švácha, P., and J. F. Lawrence, 2.1 Vesperidae Mulsant, 1839; 2.2 Oxypeltidae Lacordaire, 1868; 2.3 Disteniidae J. Thomson, 1861; 2.4 Cerambycidae Latreille, 1802, In Handbook of zoology, Arthropoda: Insecta; Coleoptera, beetles, Volume 3: Morphology and systematics (Phytophaga), eds. R. A. B. Leschen and R. G. Beutel, Walter de Gruyter, Berlin, 2014, 16–177.) D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-212.jpg&w=239&h=149 41General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.67 Coccoderus sexmaculatus Buquet (Cerambycinae). FIGURE 1.68 Compsibidion divisum Martins (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-213.jpg&w=235&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-214.jpg&w=155&h=269 42 Cerambycidae of the World FIGURE 1.69 Compsocerus deceptor Napp (Cerambycinae). FIGURE 1.70 Disaulax hirsuticornis (Kirby) (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-215.jpg&w=191&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-216.jpg&w=183&h=269 43General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.71 Lissonotus spadiceus Dalman (Cerambycinae). FIGURE 1.72 Megacyllene (Megacyllene) patruelis (Chevrolat) (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-217.jpg&w=166&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-218.jpg&w=131&h=269 44 Cerambycidae of the World FIGURE 1.73 Mionochroma chloe (Gounelle) (Cerambycinae). FIGURE 1.74 Neoregostoma coccineum (Gory) (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-219.jpg&w=121&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-220.jpg&w=128&h=269 45General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.75 Pronuba decora Thomson (Cerambycinae). FIGURE 1.76 Psygmatocerus wagleri Perty (Cerambycinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-221.jpg&w=130&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-222.jpg&w=204&h=269 46 Cerambycidae of the World 1.4.1.2 Comments Due to the diversity of forms, the subfamily is one of the most dif�cult to de�ne, with uncertain limits and relationships (Napp, 1994). Two groups of genera previously included in the Necydalinae are recognized as incertae sedis of Cerambycinae by Švácha and Lawrence (2014): (1) Atelopteryx Lacordaire, Callisphyris Newman, Hephaestion Newman, Parahephaestion Melzer, Planopus Bosq and Stenorhopalus Blanchard; and (2) Cauarana Lane, Mendesina Lane, Rhathymoscelis Thomson and Hephaestioides Zajciw. 1.4.1.3 Diversity and Distribution This is the second largest subfamily in Cerambycidae, with 1,757 genera and more than 11,200 species in the world (Bouchard et al. 2011; Monné 2012; Tavakilian 2015) (Table 1.1). The Cerambycinae are widely distributed in all biogeographic regions. In the Australian, Nearctic, and southern Neotropical regions, the Cerambycinae are the most speciose subfamily compared to other subfamilies (e.g., Forchhammer and Wang 1987; Švácha and Lawrence 2014). 1.4.1.4 Biology Adults are extremely diverse, from dark nocturnal forms to brightly colored mimetic diurnal species (Švácha and Lawrence 2014) (Figures 1.67 through 1.76). Linsley (1962b) has attempted to divide this subfamily into two groups based on adult activity patterns. Adults of many species from the Callidiopini, Gracilliini, Opsimini, Methiini, and so on are active during the night, for example, Oemona hirta (F.) (Wang et al. 1998). Some adults may be crepuscular (vespertine) such as Nadezhdiella cantori (Hope) (Wang et al. 2002), Phoracantha semipunctata (F.), and P. recurva Newman (Wang at el. 2008). Nocturnal or crepuscular adults may or may not need to feed depending on species. Adults of most species appear to be diurnal, such as spe- cies from the Aphneopini, Callidiini, Cleomenini, Clytini, Molorchini, and so forth. Many diurnally active adults visit �owers and feed on pollens and nectar, such as Zorion guttigerum Westwood (Wang 2002). Some cerambycine adults feed on tree foliage, such as Lissonotus spadiceus Dalman (Figure 1.71) (M. L. Monné, personal observation) and Xylotrechus pyrrhoderus Bates (Guo 1999); a few in Trachyderini feed on fruit. Adults can live for a week to a few months depending on whether they feed. Male-produced long-range sex and aggregation pheromones have been identi�ed in many cerambycine species (see Chapter 5). Mating may occur on larval hosts or adult feeding sites (see Chapter 4). Depending on the nature of larval feeding biol- ogy, adults may be attracted to larval hosts of certain conditions for oviposition (Hanks 1999; see Chapter 3). They then lay their eggs on the surface of larval host plants or in crevices and wounds of bark or under loose bark. Each female can lay dozens to hundreds of eggs in her lifetime. The distinctive cerambycine larval mouthparts are well suited for solid hosts; most larvae do not occur in soft rotten wood or in soil, and species feeding in soft herbs are rare (Švácha and Lawrence 2014). Although larvae of many species feed on dead (not rotten) plants, most species probably attack living plants that may be perfectly healthy, weakened, or stressed (Hanks 1999). The larval host range of cerambycine species can be from oligophagous to highly polyphagous across both angiosperms and gymnosperms (see Chapters 11 and 12). Larvae bore in branches and stems of host plants and sometimes enter roots, such as O. hirta (Wang et al. 1998). Mature larvae usually pupate in their host plants. The life cycle usually lasts one to four years. Many cerambycine species are important pests of trees and logs. 1.4.2 Subfamily Dorcasominae Lacordaire, 1868 1.4.2.1 Diagnosis Small to moderately large, usually elongate beetles with tapering or subparallel elytra and often long cursorial legs; no strongly depressed forms. Head prognathous or distinctly rostrate, usually constricted immediately behind eyes but never with prominent temples followed by a constricted neck; antennal insertions of variable position but at least slightly away from mandibular condyle, antennal sockets D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 47General Morphology, Classi�cation, and Biology of Cerambycidae usually facing laterally or laterodorsally; eyes moderate to very large, emarginate to entire; mandibles never enlarged, apex unidentate, inner margin usually with a distinct fringe of hairs; maxillae and labium well developed, lacinia distinct, submentum with a very short to long intermaxillary process, ligula usu- ally large, membranous and emarginate or bilobed, terminal segments of both palps usually more or less truncate. Pronotum without lateral carina, often with a pair of lateral tubercles or spines; procoxal cavities closed internally and at least narrowly open posteriorly, prosternal process usually narrow but complete; notosternal suture may be relatively distinct and complete; mesonotum usually with a divided stridulatory plate; mesocoxal cavities open laterally. Elytra in some taxa strongly narrowed and separate or also shortened posteriorly, partly exposing hind wings yet almost always distinctly surpassing poste- rior pterothoracic margin; hind wing with radial cell closed proximally but without edge cell. Legs short to moderately long; procoxae transverse to subglobular, prominent, projecting at least slightly below prosternal process; tarsi pseudotetramerous and padded beneath, tarsomere 5 in males of some taxa remarkably broadened distally, claws free, divaricate to moderately divergent. Empodium indistinct. 1.4.2.2 Comments Some authors treat this group of cerambycids as two separate subfamilies, Dorcasominae and Apatophyseinae, each with one tribe (e.g., Danilevsky 1979; Tavakilian 2015). However, most authors accept that these two subfamilies should be two tribes, Apatophyseini and Dorcasomini, under the sub- family Dorcasominae (e.g., Švácha and Danilevsky 1987, 1989; Özdikmen 2008; Švácha and Lawrence 2014; Adlbauer et al. 2015; Vives 2015). We adopt the latter opinion in this chapter. 1.4.2.3 Diversity and Distribution There are about 340 described species in 95 genera and two tribes occurring in the Oriental, southern Palaearctic, and Afrotropical regions (Švácha and Lawrence 2014; Adlbauer et al. 2015; Tavakilian 2015; Vives 2015) (Table 1.1). 1.4.2.4 Biology Similar to the Lepturinae, adults of many dorcasomine species are diurnal with some apatophyseine species being �oricolous (Švácha and Lawrence 2014). Most adults may be nocturnally active and hide under the bark of trees or between dead logs and the ground. For example, the nocturnal Apterotoxitiades vivesi Adlbauer adults are found under one- to two-year-old pine logs lying on the ground adjacent to grassland (Adlbauer et al. 2015). It is not clear whether adults feed and how they reproduce. No long- range pheromones have been found. Švácha and Lawrence (2014) summarized the known biology of this subfamily. Larvae of Dorcasomus gigas Aurivillius make wide galleries along the center of stems and branches of living trees and pupate in the host plant. Larvae of Apatophysis Chevrolat develop in dead or moribund underground parts of trees and shrubs and in dry, often treeless, habitats with large perennial herbs. Adlbauer et al. (2015) speculate that A. vivesi larvae may feed on grass roots. Undescribed larvae of many Madagascan and one South African (Otteissa Pascoe) genera were found in dead, often rotting, wood, mostly above the ground; but some species are subterranean (and larvae also tend to lose stemmata). They are found less frequently in relatively fresh dead branches where larvae usually feed subcortically; unidenti�ed dorca- somine larvae were also found in the outer bark layer of large living broad-leaved trees. Mature larvae of nearly all known species leave the host material and pupate in soil. The life-cycle length is unknown for this subfamily. 1.4.3 Subfamily Lamiinae Latreille, 1825 1.4.3.1 Diagnosis Small to large, elongate to robust beetles (Figures 1.77 through 1.91). Head (Figure 1.7) vertical in front or retracted and hypognathous, genal line directly posterior; antennae inserted high on frons between eyes; D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 48 Cerambycidae of the World FIGURE 1.77 Demophoo hammatus (Chabrillac) (Lamiinae). FIGURE 1.78 Hamastatus conspectus Monné (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-223.jpg&w=198&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-224.jpg&w=202&h=269 49General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.79 Hydraschema fabulosa Thomson (Lamiinae). FIGURE 1.80 Hylettus stigmosus Monné (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-225.jpg&w=104&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-226.jpg&w=191&h=269 50 Cerambycidae of the World FIGURE 1.81 Lycaneptia nigrobasalis Tippmann (Lamiinae). FIGURE 1.82 Macronemus �licornis (Thomson) (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-227.jpg&w=182&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-228.jpg&w=168&h=269 51General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.83 Megabasis speculifera (Kirby) (Lamiinae). FIGURE 1.84 Onychocerus albitarsis Pascoe (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-229.jpg&w=149&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-230.jpg&w=159&h=269 52 Cerambycidae of the World FIGURE 1.85 Pertyia sericea (Perty) (Lamiinae). FIGURE 1.86 Melzerella lutzi Lima (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-231.jpg&w=188&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-232.jpg&w=145&h=269 53General Morphology, Classi�cation, and Biology of Cerambycidae FIGURE 1.87 Oncideres ulcerosa (Germar) (Lamiinae). FIGURE 1.88 Onocephala obliquata Lacordaire (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-233.jpg&w=147&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-234.jpg&w=170&h=269 54 Cerambycidae of the World FIGURE 1.89 Psapharochrus luctuosus (Bates) (Lamiinae). FIGURE 1.90 Scleronotus scabrosus Thomson (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-235.jpg&w=165&h=269 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-236.jpg&w=162&h=269 55General Morphology, Classi�cation, and Biology of Cerambycidae second antennomere small; eyes �nely or coarsely faceted, emarginate, frequently divided; labrum free; mandibles acute without a molar plate and incisor edge without fringe of hairs; maxillae and labium well-developed; palpi with last segment pointed at apex (Figures 1.14 and 1.15); ligula membranous; mentum trapezoidal; gula usually with a short intermaxillary mentigerous process. Pronotum without a lateral margin; procoxae globose or subconical, cavities usually angulated externally (Figure 1.26); mesoscutum usually with a complete endocarina and an undivided stridulatory plate (Figure 1.32); meso- coxal cavities usually open to epimera. Legs usually short; protibiae with an oblique internal mesial sinus; mesotibiae usually with an oblique external sinus, tibial spurs short; tarsi pseudotetramerous, third tarsomere dilated, concealing the minute fourth tarsomere; tarsal claws divaricate, divergent, appendicu- late, or bi�d; empodium absent. 1.4.3.2 Comments The subfamily Lamiinae had always been considered a very well-de�ned and monophyletic group (Napp 1994; Švácha and Lawrence 2014). Audinet-Serville (1832) was the �rst to propose a classi�cation of the Cerambycidae, which included three “tribus”: Prioniens, Cerambycins, and Lamiares. Audinet-Serville (1834) divided Lamiares by the body form into “Déprimés” and “Convexes,” describing numerous gen- era and species. This classi�cation generally was accepted in Europe by Bates (1861, 1862, 1863, 1864, 1865, 1866), Pascoe (1864–1869), and Thomson (1860, 1861, 1864). LeConte and Horn (1883) recognized the Lamiinae by the prothorax not being margined, the palpi with the last joint cylindrical and pointed, and frequently the front tibiae obliquely sulcate on the inner side. They arranged the tribes represented in North America into nine series (including “Lamioides”) and several subdivisions. Linsley and Chemsak (1984) were unable to resolve contradictions and inconsistencies in the recent usage of Lamiinae tribal classi�cation, mainly in North America. The classi�cation proposed by Linsley and Chemsak represents a compromise between that of LeConte and Horn and more recent work, thus FIGURE 1.91 Trachysomus fragifer (Kirby) (Lamiinae). D ow nl oa de d by [ M as se y U ni ve rs ity L ib ra ry ], [ Q ia o W an g] a t 1 7: 05 0 9 M ay 2 01 7 http://www.crcnetbase.com/action/showImage?doi=10.1201/9781315313252-2&iName=master.img-237.jpg&w=147&h=269 56 Cerambycidae of the World retaining as many of the familiar names as possible of those currently in use by researchers of North American longicorn fauna. 1.4.3.3 Diversity and Distribution This is the largest subfamily, with more than 21,000 species in 2,964 genera. It has representatives in all biogeographic regions and is particularly diverse in the tropics and subtropics (Table 1.1). Although the Lamiinae are the species-richest subfamily in most regions, they are outnumbered by the Cerambycinae in the Australian, Nearctic, and Neotropical regions (Forchhammer and Wang 1987). 1.4.3.4 Biology A number of authors have summarized the general biology of the Lamiinae (e.g., Butovitsch 1939; Linsley 1961; Linsley and Chemsak 1984, 1995; Wang and Chiang 1991; Hanks 1999; Wang 2008; Ślipiński and Escalona 2013; Švácha and Lawrence 2014). Adults of most lamiine species probably are nocturnal or crepuscular in habit, based on evidence from morphological (such as coarsely faceted eyes) and biological studies. Many are diurnal, and a few are �ower visitors. In almost all species of known biology, adults feed before sexual maturation; for example, many species feed for an average of seven days before mating (Hanks 1999). Most adults feed on leaves and stems of their larval hosts (Hanks 1999), such as Paraglenea fortunei Saunders (Wang et al. 1990), Phytoecia ru�ventris Gautier (Wang et al. 1992), and Glenea cantor (F.) (Lu et al. 2011). Others feed on conifer needles and cones, such as Acanthocinus and Monochamus species, and a few on pollen, stamens, nectar, fungus, or fermented tree oozes (Butovitsch 1939). Due to the nature of adult feeding, active adult life span is relatively long, rang- ing from weeks to months. Flightlessness is not infrequent and obviously of multiple origin and appar- ently always occurs in both sexes. So far, only male-produced long-range aggregation pheromones have been identi�ed for lamiines (see Chapter 5). Lamiine adults usually prepare the oviposition sites in the bark of stems or branche