Handbook of Zoology Founded by Willy Kükenthal Editor-in-chief Andreas Schmidt-Rhaesa Arthropoda: Insecta Editors Niels P. Kristensen & Rolf G. Beutel Authenticated | [email protected] Download Date | 5/8/14 6:22 PM Richard A. B. Leschen Rolf G. Beutel (Volume Editors) Coleoptera, Beetles Volume 3: Morphology and Systematics (Phytophaga) Authenticated | [email protected] Download Date | 5/8/14 6:22 PM Scientific Editors Richard A. B. Leschen Landcare Research, New Zealand Arthropod Collection Private Bag 92170 1142 Auckland, New Zealand Rolf G. Beutel Friedrich-Schiller-University Jena Institute of Zoological Systematics and Evolutionary Biology 07743 Jena, Germany ISBN 978-3-11-027370-0 e-ISBN 978-3-11-027446-2 ISSN 2193-4231 Library of Congress Cataloging-in-Publication Data A CIP catalogue record for this book is available from the Library of Congress. Bibliografic information published by the Deutsche Nationalbibliothek The Deutsche Nationalbibliothek lists this publication in the Deutsche Nationalbibliografie; detailed bibliographic data are available in the Internet at http://dnb.dnb.de Copyright 2014 by Walter de Gruyter GmbH, Berlin/Boston Typesetting: Compuscript Ltd., Shannon, Ireland Printing and Binding: Hubert & Co. GmbH & Co. KG, Göttingen Printed in Germany www.degruyter.com Authenticated | [email protected] Download Date | 5/8/14 6:22 PM Cerambycidae Latreille, 1802 77 2.4 Cerambycidae Latreille, Batesian mimic (Elytroleptus Dugés, Cerambyc inae) feeding upon its lycid model (Eisner et al. 1962), 1802 the wounds inflicted by the cerambycids are often non-lethal, and Elytroleptus apparently is not unpal- Petr Svacha and John F. Lawrence atable or distasteful even if much of the lycid prey is consumed (Eisner et al. 2008). In some cases, Distribution. A worldwide family with approxi- for instance the East Asian lamiine genus Doliops mately 35,000 described species (database Titan). Waterhouse mimicking various species of the cur- The family is presently divided into eight sub- culionid genus Pachyrhynchus Germar, the model is families: Prion inae (over 1000 species), Parandr- not known to be noxious. However, unlike Doliops, inae (119), Dorcasom inae (over 300), Cerambyc inae the weevils are extremely heavily sclerotized and (ca. 11,000), Spondylid inae (ca. 100), Necydalinae may be mechanically protected. In some species (ca. 70), Lepturinae (ca. 1500), and Lamiinae (over possessing very different discrete color forms and/ 20,000). Species richness is highest in the tropics, or remarkable sexual dimorphism/dichroism (e.g., where the fauna comprises mainly taxa belong- some anacoline Prioninae), either one form or sex ing to the subfamilies Prioninae, Cerambyc inae may be mimetic, or each possibly mimics a differ- and Lamiinae. Some higher taxa (at the level of ent model; in extreme cases, the male and female subfamilies, the primarily Northern Hemisphere beetles are difficult to reconcile as conspecifics Necydalinae, Lepturinae and Spondylidinae) are (Fig. 2.4.2 L, M). absent or scarce in the tropics and often limited Bright coloration of some cerambycids is con- to higher elevations. Dorcasominae is very diverse sidered true aposematism. The lamiine genus (second largest following Lamiinae) in Madagas- Tetraopes Dalman in Schoenherr, with predomi- car. Only four subfamilies (Prion inae, Parandr inae, nantly red adults, sequesters cardenolides from Cerambyc inae, and Lamiinae) occur in Australia, its host-plant Asclepias, although the effectiveness New Zealand and the Pacific islands, and Australia of the chemical protection has been questioned belongs to the very few major regions where species (references in Allison et al. 2004), and Linsley of Lamiinae are outnumbered by Cerambycinae (1959) cites examples of birds feeding on Tetraopes (McKeown 1947; Forchhammer & Wang 1987). beetles. Other antipredatory modes, such as iri- descent colors that abruptly change with viewing Biology and Ecology. Adult defense and mime- angle or brightly colored abdomens visible only in tism. Ancestral cerambycids were probably dull, flight, are common in cerambycids. The lamiine somber-colored, crepuscular or nocturnal beetles. Onychocerus albitarsis Pascoe has spine-like termi- Such species still prevail in Prion inae, Parandr- nal antennomeres (present also in other species) inae, Spondylid inae, and in the two related fami- modified as scorpion-like stingers that are used in lies Disteniidae and Vesperidae. Some nocturnal defense (Berkov et al. 2008). adults are hidden during the day (they may even Both sexes in subfamilies other than Prioninae return to their exit galleries), and their adaptations and Parandr inae usually possess a stridulatory are generally “mechanical”, such as antipredatory device consisting of a striated plate on the meso- spines or pilosity, burrowing modifications, etc. scutum (Fig. 2.4.14 D–G) and ridge(s) on the ven- However, the perplexing diversity of color and tral face of the posterior pronotal margin. Because form and the clear mimetism of many forms active both sexes usually stridulate when disturbed or or exposed during the day suggest that visually handled, the sound is assumed to be defensive, orienting vertebrate predators are high on the list though some adults also produce sounds during of their enemies. Many cerambycids (particularly courtship and copulation. Certain Prion inae devel- Lamiinae) are cryptic, resembling bark, lichens or oped a different sound-producing mechanism as a even bird droppings. Although crypsis is useful defense (rubbing ridged hind femora against finely to both diurnal and nocturnal species, mimicry striate lateral elytral margin). occurs more in day-active forms. Lycid, cantharid Adult feeding. Butovitsch (1939) attempted to or meloid beetles (Fig. 2.4.4 F) or aculeate Hymeno- classify the types of adult feeding in Cerambyc idae. ptera (Fig. 2.4.4 U, 2.4.5 J, N, 2.4.6 K, L, 2.4.8 G) In many presumably basal groups (Parandrinae, are models for Necydalinae and some members of most Prion inae, many Cerambyc inae, Spondylid- the remaining subfamilies, except Parandr inae. inae), the adults do not feed and the midgut may be Hymenopteran mimicry may involve body shape, rudimentary and thread-like, or they may imbibe color pattern, reduced elytra and exposed hind fluids, such as fermenting sap, or feed on ripe or wings (sometimes without apical folding) and fermenting fruits. Samuelson (1994) proposed pol- characteristic movements. Ant mimics may occur linophagy as the ancestral type of chrysomeloid among their models on the host trees (Vives et al. adult feeding, but this proposal does not fit ceram- 2011; Vives 2012) but inquilines are unknown. bycoids, in which presumed basal forms are often Cerambycid mimicry has been mostly assumed to large and/or lack a suitable type of mouthparts. be Batesian, although palatability of mimics was Widespread floricoly in the Lepturinae, Dorcasom- seldom rigorously tested. In a very unusual case of a inae and Cerambyc inae does not extend to some of Authenticated | [email protected] Download Date | 5/8/14 6:21 PM 78 Petr Svacha and John F. Lawrence their possible basal groups; however, feeding on and relatives, Callidiini, Clytini or Curiini: Nearns anemophilous pollen, spores or similar material & Ray 2006; Ray et al. 2006). The larger, more com- might be ancestral and precede modified mouth- plex prothorax in many males of various tribes parts with a prominent galea and lacinia bearing (particularly in Cerambyc inae and Prion inae) long, often apically curved, setae. Crepuscular flo- may be associated with production of such phero- ricoly has been recorded by Danilevsky & Mirosh- mones (Fig. 2.4.14 A–C). Male-produced volatile nikov (1981) for the lepturine Enoploderes sanguineum pheromones cannot be expected (and associated Faldermann that visits flowers of Swida sanguinea at modified male prothoraces do not occur) in some sunset. The universal obligatory adult feeding of Prion inae in which females are flightless, at least Lamiinae on living plants (typically leaves or fresh until a portion of the eggs are laid (virtually no bark, usually but not always of the same plant taxon Cerambyc inae have flying males and flightless that serves as larval host) or on dead bark and fungi females). A short-range, female-produced vola- is undoubtedly apomorphic. Fungal fruiting bod- tile sex pheromone was implied in Semanotus ies are a poorly known but possibly overlooked and japonicus (Lacordaire) by Fauziah et al. (1992). relatively widespread adult food source for certain True long-range female sex pheromones were lamiine taxa (review and references in Adlbauer presumed or behaviorally demonstrated in some 2004), and records of other Lamiinae feeding on Prion inae (Prionoplus reticularis White: Edwards bark of dead branches (e.g., some Mesosa Latreille or 1961 b; Prionus californicus Motschulsky: Cervantes Pogonocherus Dejean in Cherepanov 1983, 1984) may et al. 2006; Barbour et al. 2006) and Cerambyc- in part concern unrecognized fungivory. Pollen- or inae (Callisphyris Newman: Krahmer 1990). The nectar-feeding is very rare in Lamiinae and prob- pheromone of P. californicus is produced in glands ably just a supplementary food source. Adults of associated with the ovipositor and was identified an undescribed Chinese lamiine species of Falsome- as (3R,5S)-3,5-dimethyldodecanoic acid (Rodstein sosella Pic that developed in fallen rotting branches et al. 2009, 2011). In Lepturinae, (Z)-11-octadecen- of a broadleaved tree produced dark feces while in 1-yl acetate was identified as a probable female pupal chambers and are thought to have fed on the long-range pheromone in Ortholeptura Casey (Ray (possibly fungal) material on the walls of the pupal et al. 2011), and (4R,9Z)-hexadec-9-en-4-olide in cells (C. Holzschuh, personal communication). Desmocerus Dejean (Ray et al. 2012). Because long- Whereas the life span of emerged adults is usu- range pheromones require high sensitivity by the ally measured in weeks or even days for many receiver, males often have flattened, serrate or pec- species, certain larger Lamiinae may be active for tinate/flabellate antennae with large sensory sur- several months under laboratory conditions when faces (Fig.