The Book of A Life-Size Guide to Six Hundred of Nature's Gems

Edited by Patrice Bouchard

ith 350,000 known This collection covers six hundred , and scientific significant species. Each W estimates that mil- features a distribution map, basic lions more have yet to be identi- , conservation status, and fied, beetles are one of the most information on cultural and eco- remarkable and varied creatures nomic significance. Full-color on earth. They range from the de- photos show the beetles both at lightful summer firefly to the one- actual size and enlarged to show hundred-gram Goliath beetle. details. Based in the most up-to- Beetles offer a dazzling array of date science and accessibly writ- shapes, sizes, and colors that en- ten, the descriptive text will ap- tice scientists and collectors across peal to researchers and armchair the globe. coleopterists alike.

Contributors:

PATRICE BOUCHARD, YVES BOUSQUET, CHRISTOPHER CARLTON, MARIA LOURDES CHAMORRO, HERMES E. ESCALONA, ARTHUR V. EVANS, ALEXANDER S. KONSTANTINOV, RICHARD A. B. LESCHEN, STÉPHANE LE TIRANT, AND STEVEN W. LINGAFELTER.

Publication date For a review copy or other To place orders in the October 15, 2014 publicity inquiries, please United States or Canada, contact: please contact your local $55.00, cloth Lauren Salas University of Chicago Press 978-0-226-08275-2 Promotions Manager, sales representative or contact the University of 656 pages University of Chicago Press, Chicago Press by phone at 2400 color plates [email protected] 773-702-0890 1-800-621-2736. 7 1/8 x 10 1/2

THE BOOK OF BEETLES THE BOOK OF BEETLES

A LIFE-SIZE GUIDE TO SIX HUNDRED OF NATURE’S GEMS

EDITOR PATRICE BOUCHARD

CONTRIBUTORS PATRICE BOUCHARD, YVES BOUSQUET, CHRISTOPHER CARLTON, MARIA LOURDES CHAMORRO, HERMES E. ESCALONA, ARTHUR V. EVANS, ALEXANDER KONSTANTINOV, RICHARD A. B. LESCHEN, STÉ PHANE LE TIRANT, STEVEN W. LINGAFELTER

THE UNIVERSITY OF CHICAGO PRESS Chicago PATRICE BOUCHARD is a research scientist and curator of Coleoptera at the Canadian National Collection of , Arachnids, and Nematodes. CONTENTS YVES BOUSQUET is a research scientist at the Canadian National Collection of Insects, Arachnids, and Nematodes. CHRISTOPHER CARLTON is a research scientist and director of the Louisiana State Museum MARIA LOURDES CHAMORRO is a research entomologist with the Systematic Entomology Laboratory at Introduction 6 the National Museum of Natural History in Washington, D.C. HERMES E. ESCALONA is a visiting scientist at the Australian National Collection-CSIRO. ARTHUR V. EVANS is a research associate at the Smithsonian Institution. What is a beetle? 10 ALEXANDER KONSTANTINOV is a research entomologist with the Systematic Entomology Laboratory at the National Museum of Natural History in Washington, D.C. Beetle classi cation 16 RICHARD A. B. LESCHEN is a researcher at Landcare Research, New Zealand Arthropod Collection. Evolution & diversity STÉPHANE LE TIRANT is curator of the Montreal Insectarium. 18 STEVEN W. LINGAFELTER is a research entomologist with the Systematic Entomology Laboratory, Communication, reproduction & development 20 United States Department of Agriculture Defense 22

The University of Chicago Press, Chicago 60637 This book was conceived, Feeding behavior 24 The University of Chicago Press, Ltd., London designed, and produced by © 2014 by Ivy Press Limited Ivy Press Beetle conservation 26 All rights reserved. Published 2014. 210 High Street, Lewes Printed in China East Sussex BN7 2NS United Kingdom Beetles & society 28 23 22 21 20 19 18 17 16 15 14 1 2 3 4 5 www.ivy-group.co.uk

ISBN-13: 978-0-226-08275-2 (cloth) Creative Director PETER BRIDGEWATER The beetles 30 ISBN-13: 978-0-226-08289-9 (e-book) Publisher SUSAN KELLY DOI: 10.7208/chicago/9780226082899.001.0001 Art Director MICHAEL WHITEHEAD 32 Editorial Director TOM KITCH 42 Portions of this work were written and prepared by o cers and/ Senior Project Editor CAROLINE EARLE or employees of the U.S. Government as part of their o cial duties Commissioning Editor KATE SHANAHAN 48 and are not copyrightable. Designer GINNY ZEAL 112 Illustrator SANDRA POND Library of Congress Cataloging-in-Publication Data LITHOCASE IMAGES Appendices 640 The book of beetles : a life-size guide to 600 of nature’s © Jason Bond and Trip Lamb: Onymacris bicolor. gems / edited by Patrice Bouchard ; with contributions by © Lech Borowiec: aurata, Dytiscus marginalis, Lytta vesicatoria, Glossary 642 Patrice Bouchard, Yves Bousquet, Christopher Carlton, Pachylister inaequalis, Pachnephorus tessellates Classi cation of the Coleoptera 646 Maria Lourdes Chamorro, Hermes E. Escalona, Arthur V. Karolyn Darrow © The Smithsonian Institution: Tetracha carolina Evans, Alexander Konstantinov, Richard A. B. Leschen, Resources 648 Stéphane Le Tirant, and Steven W. Lingafelter. Anthony Davies © Her Majesty the Queen in Right of Canada as represented by the Minister of Agriculture and Agri-Food: Actinus pages : illustrations ; cm Notes on contributors 650 imperialis, museorum, Brachycerus ornatus, Byctiscus rugosus, Includes bibliographical references and index. Erotylus onagga, Hister quadrinotatus quadrinotatus, Saprinus cyaneus, Index of species and families 652 ISBN 978-0-226-08275-2 (cloth : alk. paper) — ISBN Spilopyra sumptuosa 978-0-226-08289-9 (e-book) 1. Beetles—Identification. Acknowledgments 656 2. Beetles—Pictorial works. I. Bouchard, Patrice, 1973– Henri Goulet © Her Majesty the Queen in Right of Canada as represented by the Minister of Agriculture and Agri-Food: sycophanta editor. QL575.B66 2014 © René Limoges: atlas 595.76—dc23 © Kirill Makarov: Byturus tomentosus 2014010716 © Udo Schmidt: Paranaleptes reticulata ∞ This paper meets the requirements of ANSI/NISO Z39.48- © Maxim Smirnov: Lamprima adolphinae, Mormolyce phyllodes, 1992 (Permanence of Paper). Sulcophaneus imperator © Chris Wirth: Alaus zunianus, Amblysterna natalensis, Calodema regalis, Color origination by Ivy Press Reprographics Thrincopyge alacris

[CHECK UCP PLC IMAGES] INTRODUCTION

or hardened forewings, or elytra (singular elytron). Depending on the species, elytra can help stabilize beetles in flight, protect their delicate hind wings and internal organs, conserve precious bodily fluids, capture bubbles of air underwater, and insulate them from extreme temperatures. Combined with their small and compact bodies and numerous other morphological and behavioral adaptations, beetles exploit and thrive in niches unoccupied or underutilized by other in widely diverse 6 INTRODUCTION terrestrial and freshwater habitats. Although the sheer number of species prevents all but the most common or economically important beetles from having a meaningful and widely accepted common name, each known species does have a scienti c name consisting of

a (plural genera) and species (singular and plural) epithet that is ABOVE Some beetles feed on a single plant universally recognized. To manage information eectively, coleopterists species; others feed on a wide range of file each species into a nested system of hierarchical groups, or taxa hosts. Adults of the North American (singular taxon), based on their shared evolutionary characteristics. the Japanese Beetle (Popillia japonica) Species is the most exclusive taxon, while the order Coleoptera is the most have been observed feeding on 300 hosts inclusive of beetle taxa. in approximately 80 plant families. “From the small size of insects, we are apt to undervalue their appearance. Beetles communicate with one another through physical, chemical, or visual means, usually to locate a mate. Although most species engage in BELOW Most beetles If we could imagine a male Chalcosoma with its polished, bronzed coat of are winged, but some sexual reproduction, a few reproduce asexually by cloning themselves, a have reduced hind mail, and vast complex horns, magni ed to the size of a horse or even of a dog, wings and cannot y, process known as parthenogenesis. Among beetles, limited parental care such as species in the it would be one of the most imposing animals in the world.” African genus of the young is the exception, not the rule. The larvae and adults eat a Brachycerus. CHARLES DARWIN, THE DESCENT OF MAN, & SELECTION IN RELATION TO SEX, 1871 of organisms, living and dead, especially plants. Those that prefer leaves, owers, fruits, needles, cones, and roots can

ABOVE Beetles Beetles of the order Coleoptera, with nearly 400,000 described species, inict serious damage to food stores, gardens, crops, and represent an exceptionally diverse comprise one of the most diverse and important groups of animals on managed timber. Some carnivorous beetles are used group of organisms. Their importance for Earth. As such, coleopterists, biologists who specialize in the study of as biological control agents against agricultural or agriculture, forestry, culture, and science, beetles, have a view of the natural world with a degree of resolution that forestry pests, while scavenger species provide and endless variations in structures (e.g., the is seldom seen through the study of other organisms. an essential service to clean study skeletons male , Chalcosoma atlas) One out of every ve species of plants and animals is a beetle. Despite in natural history collections around the and microhabitat adaptations, have their riot of forms, colors, patterns, and behaviors, all beetles share a select globe. Recently, the study of beetle structure captivated humans for centuries. suite of physical attributes, the most conspicuous of which are the leathery and function has inspired scientists and engineers INTRODUCTION

Physically impressive Large, colorful, horned, or exaggerated or unusually developed legs or mouthparts—species with special structures involved in speci c behavior that has evolved through natural selection over millions of years and thus carries interesting underlying genetic information.

BELOW Millions of Each species is depicted by a razor-sharp life-size photograph and insects, including beetles, are preserved accompanied by a summary of its known distribution, habitat, and feeding in natural history collections throughout habits. The map oers an indication of its global distribution, while the the world. Specimens are typically dried and engravings afford readers another point of view. The scientific and pinned, with a label common (if any) names are also provided, followed by the species’ author attached to the same pin that includes and year of description. Concise narratives covering natural history and important information such as the locality related species are followed by a brief species diagnosis. where the individual 8 was encountered, the 9 date it was collected, and the person who BEETLE COLLECTIONS found it. Specimens are then placed in Beetle collections carefully assembled over decades by thousands of boxes or drawers, such as this one from dedicated professionals and amateurs provide essential data needed to the Alfred Russel Wallace (1823–1913) identify and map sensitive species within a historical context. These collection in the Natural History ABOVE One of the most working in the rapidly growing eld of biomimetics to develop and design scienti cally valuable collections also serve as important resources for Museum, London, in challenging tasks order to protect them for this project was new materials and products ranging from iridescent car paint and reusable other scienti c and educational endeavors, such as providing the specimens from museum pests to select the 600 and improve long- candidate beetles adhesive tape to monetary security systems. photographed for this book. term preservation. to cover from the overwhelming The Book of Beetles oers a glimpse of this incredible diversity through number of species. While some species an overview of 600 beetle species presented within a framework based on such as the Agreeable Caterpillar Hunter their evolutionary relationships. The diversity of beetles is divided by (Calosoma sycophanta) (left) are visually species into four main chapters of the suborders. Within each section the attractive and well known biologically, arrangement is taxonomic, by family and then , and then within others such as the Frog-legged Leaf each subfamily in scienti c name (genus, species) order. Beetle (Sagra buqueti) (right) have notable unusual adaptations yet little is known SELECTION CRITERIA about their biology. We hope that this The species were selected on the basis of several criteria, oering a unique book will stimulate the discovery and taxonomic survey of the majority of beetle families from around the world: publication of new knowledge on beetles Scienti cally compelling Subjects of focused scienti c research or medicinal from around the world. use, or inspirations for biomimetic and technological innovation. Curious natural histories Unusual adaptations, ability to live in extreme habitats, interesting symbioses, or engaging behaviors. Culturally signi cant Mythological and religious symbols, uses in folk medicine, or entomophagy. Economically important Pest species, use as biological control agents, sources of products and services, or with a role in forensic entomology. Conservation Rare and threatened species. WHAT IS A BEETLE?

DORSAL VIEW VENTRAL VIEW

RIGHT The forewings of beetles are mandible uniquely modied into hardened, non- antenna labrum foldable elytra that meet together in a straight line over the body when at rest. mentum The elytra partly or compound eye completely cover the front leg abdomen and can pronotum be opened to deploy prosternal the hind wings, as in scutellum process this Lucanus cervus elytron individual. elytral tibia epipleuron middle leg coxa

femur 11 10 WHAT IS A BEETLE? hind leg tarsal trochanter claw

The English word “beetle” comes from the Middle English bityl or betyll and the Old English bitula, all of which mean “little biter.” Other elytral abdominal commonly used names, such as “weevil” and “chafer” derived from Old tarsomeres suture tarsus ventrites English and Old High German, also relate to biting. Coleoptera, rst coined by Aristotle in the fourth century bce and later adopted as an order secretions or microscopic networks of cracks (alutaceous) resembling that ABOVE The study of a superdiverse group of insects by in 1758, is derived from the Greek words koleos, of human skin. The surface is variously festooned with spines, hairlike of organisms such as beetles requires the meaning “sheath,” and pteron, or “winged,” and was inspired by the tough setae, or flattened setae called scales, and sculpted with tiny bumps close examination of a wide range of elytra of beetles. (tubercles), pit-like punctures, ridges, grooves (striae), or rows of punctures. anatomical structures in order to recognize and classify species and larger groups DISTINGUISHING FEATURES COLOR correctly. Using clear and consistent terms Among other adaptations, beetles are distinguished from other insects by The colors of beetles are derived either from chemical pigments obtained for these structures is essential to clearly their chewing mouthparts, the conversion of their forewings into hardened from their food or structural properties of the outer layers of the communicate results amongst those who elytra, their hind wings that fold lengthwise and across beneath the elytra, exoskeleton. Most beetles are black as a result of melanin deposition during are specialists and non-specialists. and their holometabolous development. Holometabolous insects pass through sclerotization, the chemical hardening process of the exoskeleton that occurs four very distinct life stages: egg, , pupa, and adult. The larvae and after emergence from the pupa, or eclosion. Microscopic surface sculpturing adults frequently dier in habits and habitat, functioning in the environment also influences beetle colors, as do patterns of setae, scales, or waxy as if they were two separate species. secretions. Black desert darkling beetles (Tenebrionidae) are sometimes Beetles, like other insects, crustaceans, arachnids, millipedes, centipedes, partially or completely covered with a white, yellow, or bluish-gray waxy and their kin with segmented exoskeletons and jointed appendages bloom that reects light and helps to keep the beetle cool. (antennae, mouthparts, legs), are classi ed in the phylum Arthropoda. The brilliant iridescent and metallic colors of beetles are created by Light and durable, the beetle exoskeleton is incredibly tough and rigid or multiple reective layers in the exoskelton and scales, or a layer of highly characteristically soft and pliable, and provides protection and support. It complex photonic crystals that reect light at dierent wavelengths to serves as a platform for important tactile and chemosensory structures create speci c metallic colors and shimmering iridescence. These structures externally, while providing an internal framework that supports muscles are determined genetically, but their nal form in individual beetles is and organs. The exoskeletal surface is smooth and shiny, or dulled by waxy determined by conditions experienced during growth and development. POLYPHAGA POLYPHAGA

FAMILY Staphylinidae FAMILY Staphylinidae SUBFAMILY Aleocharinae SUBFAMILY Scaphidiinae DISTRIBUTION Nearctic: North America, north of Mexico DISTRIBUTION Oriental: Sumatra and Sulawesi, Indonesia; Sarawak, Malaysia MACROHABITAT Forests MACROHABITAT Tropical forests MICROHABITAT Nests of formicine ants MICROHABITAT Fungus on logs and standing dead trees FEEDING HABITS Adults and larvae are fed by hosts through trophallaxis FEEDING HABITS Fungivorous NOTE A specialized nest parasite of formicine ants NOTE The neck may be twice the body length

ADULT LENGTH ADULT LENGTH Adult length: ∕–¼ in ½–¾ in (5.3–6.4 mm) (13–20 mm)

XENODUSA REFLEXA DIATELIUM WALLACEI XENODUSA REFLEXA LONG-NECKED SHINING (WALKER, 1866) 166 FUNGUS BEETLE 167 PASCOE, 1863

Adults and larvae of Xenodusa reflexa are adapted to an From the head back, this remarkable insect looks like a typical obligate association with formicine ants and spend most of member of the tribe Scaphidiini of the subfamily Scaphidiinae, Actual size their lives in ant nests. Specialized glandular setae on the adult but the extremely elongate neck in both sexes render it unique abdomen produce appeasement chemicals that are attractive among rove beetles. The biology of the species is poorly to host ants. Both adults and larvae are fed by regurgitation known, but other members of the subfamily graze on eshy or by the ants (trophallaxis). They are considered nest encrusting fungi on dead wood or other organic substrates in parasites because beetle larvae compete with ant larvae for forest habitats as both adults and larvae. Adults may be care and feeding by the ants. Ants in the genus Camponotus encountered on exposed fungal surfaces day or night. They are the most frequently documented hosts for this species, but are wary and prone to flight or dropping, requiring a slow, other species breed in Formica nests and overwinter in stealthy approach if observations or photographic eorts are Camponotus nests. to be successful.

RELATED SPECIES RELATED SPECIES Four species of the genus Xenodusa occur in USA and This species is placed in the same tribe as the enormous genus southern Canada, and an additional species occurs in Mexico. Scaphidium based on recent phylogenetic studies and general Xenodusa reflexa is the most widely distributed species. It appearance, exclusive of the outsized neck. It is not likely to be differs from species that overlap in ranges by its relatively confused with any other within its range, but some larger size ( ⁄ in, or 4.5 mm) and hairy ventral surface. African brentid have elongate forebodies comprising A widespread eastern species, X. cava, is similar, but lacks parts of the thorax and neck that are supercially similar. long hairs on the ventral surface of the body.

Xenodusa re exa is a comparatively large, The Long-necked Shining Fungus Beetle is unique mahogany brown member of the among rove beetles and perhaps all beetles in tribe . The broad, re exed pronotum possessing an extremely long neck that may be twice and unusual trichome bundles along the the length of the body in some male specimens. The abdomen are characteristic of the genus. Similar neck is longer on average in males but its length is glandular trichomes are found on various parts Actual size variable in both sexes. of the bodies of other myrmecophilous beetles, and presumably also function in producing and distributing glandular appeasement chemicals. POLYPHAGA POLYPHAGA

FAMILY Staphylinidae FAMILY Staphylinidae SUBFAMILY Staphylininae SUBFAMILY Staphylininae DISTRIBUTION Australian: New Guinea and extreme northern Australia DISTRIBUTION Neotropical: central South America MACROHABITAT Tropical forests MACROHABITAT Forests near rivers and streams MICROHABITAT Leaf litter, dung, carrion MICROHABITAT The body and nests of South American Water Rats (Nectomys squamipes) FEEDING HABITS Predatory on ies FEEDING HABITS Predatory on eas NOTE Preys on ies associated with animal carcasses NOTE Phoretic on small mammals and found in their nests feeding on eas

ADULT LENGTH ADULT LENGTH /–/ in ¼–/ in (16–22 mm) (6.5–7.5 mm)

ACTINUS IMPERIALIS AMBLYOPINODES PICEUS ACTINUS IMPERIALIS AMBLYOPINODES PICEUS FAUVEL, 1878 (BRÈTHES, 1926) 182 183

Adults of this unusually large and gaudily colored rove Members of this species and others in the tribe Staphylinini, beetle frequent smelly substrates such as carrion and subtribe Amblyopinina are unique among rove beetles in their dung that are likely to attract flies. Adults actively seek adaptations to life on the bodies and in the nests of small Actual size out and destroy predators that feed on maggots and mammals, mainly rodents. Adults of this species grasp the hair opportunistically capture adult flies. The immature stages and skin between the ears of South American Water Rats presumably share the predatory habits of the adults but details (Nectomys squamipes) and travel with the animal, and the of their biology are not well documented. One source larvae occur in water rat nests. Early entomologists thought mentions an association with cardamom, without giving these beetles were parasitic on the animals, but later researchers additional details. Adults exhibit strong sexual dimorphism of revealed that they are predators of eas and serve a benecial the mandibles and head, with males often having larger heads function for the rats by reducing ea populations in their nests. and longer mandibles than females. The flattened morphology of the beetle’s head is similar to that of other unrelated species that live on mammals. RELATED SPECIES Two similar species are known from the genus, with Actinus RELATED SPECIES macleayi from northern Australia diering mainly in having a At least five other species of the genus occur in the same less strongly punctured head and pronotum, in addition region. Identication of species is based on examination of to other minor differences. In other respects A. imperialis internal male sex structures and, to a lesser extent, the supercially resembles other large members of the staphylinid arrangement of specialized hairs on the body. Other genera of subtribe Philonthina, but few are as brightly colored. the tribe are similar and may be distinguished using available keys and by mammal host associations.

Actual size

Actinus imperialis is unusually large for a Amblyopinodes piceus is an elongate, somewhat staphylinid beetle, with a metallic green head attened, somber-colored rove beetle. The head and pronotum and brilliant metallic purple elytra. in particular is unusually attened and the The abdomen possesses a triangular orange spot mandibles are adapted to grasping mammal near the posterior end. The punctures on the head hairs. Elongate, movable hairs on the underside are diagnostic for this species. of the abdomen are unique to the genus. POLYPHAGA POLYPHAGA

FAMILY FAMILY Scarabaeidae SUBFAMILY Scarabaeinae SUBFAMILY Scarabaeinae DISTRIBUTION Palearctic: southern and central , North Africa, Middle East, portions DISTRIBUTION Neotropical: northern Argentina, southern Bolivia, western Paraguay of Asia MACROHABITAT Dry forest, dry thorn forest, and pasture MACROHABITAT Steppe, forest-steppe, and semidesert MICROHABITAT Often found in pastures in cattle-raising areas MICROHABITAT In the vicinity of accumulations of fresh dung FEEDING HABITS Adults commonly found in cattle dung FEEDING HABITS Adults strain nutrients from dung; larvae eat solid waste NOTE This is the most colorful and variable species of Sulcophanaeus NOTE Regarded as a sacred symbol by the ancient Egyptians

ADULT LENGTH ADULT LENGTH 1∕–1∕ in  /–1/ in (26–40 mm) (18–28 mm)

SCARABAEUS SACER SULCOPHANEUS IMPERATOR SACRED SCARAB BEETLE SULCOPHANEUS IMPERATOR LINNAEUS, 1758 (CHEVROLAT, 1844) 208 209

Scarabaeus sacer uses its rakelike forelegs to fashion balls from Sulcophanaeus imperator adults reach their peak of activity from fresh dung, then lays an egg inside the ball before rolling it January to March. They y during the day in search of fresh away and burying it. The grub feeds and completes its feces of humans and various domesticated animals, and can be development inside the ball. These beetles were revered in common in cattle pastures. They typically work in pairs to dig ancient Egypt as symbols of Khepri, a manifestation of the sun their nests directly beside or underneath animal droppings. god Ra, because their dung-rolling activities were considered Males push plugs of dung down the tunnel to females that symbols of the forces that moved the sun, represented by the fashion them into brood balls within the brood chamber. Dung dung ball, across the sky. The species was also associated with burial reduces the loss of pasturage due to the growth of rank rebirth, and the beetles, along with their likenesses carved in herbage, releases nutrients back to the soil, and disrupts the life Actual size stone, were frequently buried with the dead. cycles of pest organisms developing in the feces.

RELATED SPECIES RELATED SPECIES Scarabaeus contains 139 species in four subgenera that The genus Sulcophanaeus consists of 14 species, all but four of inhabit Afrotropical, Palearctic, and Oriental regions. which occur in South America. Three of the latter species These small to relatively large dung beetles have four inhabit Central America, including an endemic, while the distinct teeth across the clypeus, anterior coxae and fourth species is endemic to Jamaica. With its green, gold, and femora that are not enlarged, and lack front tarsi. Scarabaeus red color forms, Sulcophanaeus imperator is by far the most sacer is distinguished from other Palearctic species, in part, by colorful and variable species in the genus. the nely notched posterior pronotal margin, and the features of the middle and hind tibiae.

The Sacred Scarab Beetle has a broad, smooth Sulcophanaeus imperator is a large and patch and narrow groove along the posterior bulky beetle that is dull to weakly shining margin of the pronotum. The middle tibiae each black mixed with brilliant metallic have two oblique rows of short setae. The tips of greens, golds, and blues, and coppery the posterior tibiae are prolonged, forming a red re ections. Males have a distinct narrow plate beneath the tarsus. Males have backward-pointing horn on the head, a reddish fringe of setae along the inner while the females lack this armature. margins of their hind tibiae. Both males and females have front tarsi. Actual size POLYPHAGA POLYPHAGA

FAMILY Tenebrionidae FAMILY Tenebrionidae SUBFAMILY Stenochiinae SUBFAMILY Stenochiinae DISTRIBUTION Endemic to Frégate Island, Seychelles DISTRIBUTION Neotropical: from Mexico south to Bolivia MACROHABITAT Tropical forest MACROHABITAT Tropical forests MICROHABITAT Adults and larvae are most commonly associated with Pterocarpus indicus MICROHABITAT Adults live on the surface of trees or under bark, and larvae in wood FEEDING HABITS Decaying wood and bark FEEDING HABITS Immature stages probably develop in decaying wood NOTE Known only on Frégate Island, a 0.8 sq miles (2 sq km) island in NOTE One of the most diverse and colorful genera in the family Tenebrionidae the Indian Ocean

ADULT LENGTH ADULT LENGTH 1–1/ in ½–¾ in (25–30 mm) (12–19 mm)

POLPOSIPUS HERCULEANUS STRONGYLIUM AURATUM FRÉGATE ISLAND GIANT STRONGYLIUM AURATUM (LAPORTE, 1840) 492 TENEBRIONID BEETLE 493 SOLIER, 1848

The Frégate Island Giant Tenebrionid Beetle is one of 12 Based on the abundance of darkling beetles in most dry beetle species ranked Critically Endangered on the environments on the planet, many people assume that all International Union for Conservation of Nature’s Red List of species in this family are dark brown to black and are ground- Threatened Species (see also the Delta , dwellers. This is clearly not the case for species in the tribe Elaphrus viridis, and American Burying Beetle, Nicrophorus Stenochiini, which includes the colorful forest-dwelling americanus), and is restricted to Frégate Island in the genera Stronglyium and Cuphotes. Strongylium is certainly

Actual size Seychelles. The accidental introduction of Brown Rats (Rattus one of the most species-rich genera in the family, with nearly norvegicus) to the island in 1995 nearly caused the extinction of 1,000 described species and many more undescribed, especially Frégate Island Giant Tenebrionid Beetle adults are pale gray to dark brown and have broadly the flightless beetle, along with the endemic Seychelles in tropical areas. Strongylium auratum is relatively abundant in rounded elytra in dorsal view, which are covered by a small number of apically rounded and shiny Magpie-robin (Copsychus sechellarum), before the rodents Neotropical forests and can be encountered at elevations of tubercles. The elytra are completely fused along were eradicated in the early 2000s. 5,000 ft (1,500 m) and more. the midline, and ight wings are absent. The legs are relatively long and males have curved tibiae. RELATED SPECIES RELATED SPECIES Flightless tenebrionid genera were previously grouped The overwhelming diversity in Strongylium, coupled with an together on the basis of the absence of flight wings and almost complete lack of comparative studies of its species, is a their sealed elytra. Recent studies on internal major impediment to taxonomic and biological studies. New organ systems such as the defense glands and the species continue to be described every year, however, female genital tube now suggest that the older primarily based on di erences in color patterns, sculpture of classication did not reect natural groupings. Although their cuticle, and sexual characters. Although most species the monotypic genus Polposipus belongs to the diverse are elongated and have well-developed ight wings, such subfamily Stenochiinae based on internal characters, its as S. auratum, others are ightless and more convex. closest relatives have not yet been established.

Strongylium auratum is an elongated beetle with bright metallic green to reddish-purple re ections on the dorsal surface and legs. Each elytron has nine distinct rows of transverse punctures. Antennae are rather long, reaching beyond the base of the pronotum. Antennomeres are slightly wider at their apex and are speckled with small circular white sensory structures. The last two abdominal Actual size ventrites are contrasting yellow-reddish in color. POLYPHAGA POLYPHAGA

FAMILY FAMILY Attelabidae SUBFAMILY Rhynchitinae SUBFAMILY Rhynchitinae DISTRIBUTION Palearctic: China, Japan, Kazakhstan, Korea, Mongolia, DISTRIBUTION Nearctic and Neotropical: USA (Arizona, New Mexico, Texas), Mexico MACROHABITAT Forest (Chihuahua, Sonora, Oaxaca) MICROHABITAT Leaf rolls, vegetation MACROHABITAT Oak forests FEEDING HABITS Host plants include species of Malus, Pyrus, Sorbaria, and Populus MICROHABITAT Foliage, leaf litter NOTE The males are known for their wrestling behavior FEEDING HABITS Epidermal tissue of Quercus spp. NOTE The larvae are leaf-miners

ADULT LENGTH ADULT LENGTH ¼–/ in ¼ in (6.5–8 mm) (5.7–6.3 mm)

BYCTISCUS RUGOSUS EUGNAMPTUS NIGRIVENTRIS BYCTISCUS RUGOSUS EUGNAMPTUS NIGRIVENTRIS (GEBLER, 1830) (SCHAEFFER, 1905) 594 595

Ritualized contests and aggressive ghting behavior This species exhibits maternal care by individually placing are known to occur between males of species in this each egg between the epidermal layers of a dead leaf of the genus. The males extend their forelegs outward host tree (oaks, Quercus spp.). In spring, a female uses while rearing up on their mid and hind legs, grabbing her mandibles to cut an oviposition scar on a leaf that each other with their forelegs and touching their rostra. dropped to the ground the previous fall. She then The elongate tarsal setae may help enhance visual displays of oviposits a single egg into a cavity she created between this aggression. Females of the charismatic Byctiscus rugosus the upper and lower epidermis and seals the epidermal roll leaves into complicated cylindrical tubes for the reception tissue by pinching with her mandibles. The larva completes of the egg. The larva feeds within the tube, where it completes its entire development by feeding on the epidermal tissues of its development. the dead leaf. Pupation takes place in the soil and adults emerge the following spring. RELATED SPECIES Byctiscus includes 27 species from the Palearctic and Oriental RELATED SPECIES regions. Species are classi ed into two subgenera: Byctiscus There are approximately 100 species included in Eugnamptus. and Aspidobyctiscus. The tribe Byctiscini has an exclusively Eugnamptus is similar to Hemilypus, Acritorrhynchites, and Old World distribution and includes 12 genera in two Essodius, but can be distinguished by characters of the rostrum, subtribes. The related Pear Leaf-roller Weevil distance between the eyes, length of basitarsal segments, (B. betulae) is known to cause damage to grapevines, prominence of the rst abdominal suture, extent of dorsal pears, and other broadleaved trees and shrubs. punctation, and elytral length as it relates to the pygidium.

Actual size Actual size

Byctiscus rugosus is a bright metallic, shiny green weevil Eugnamptus nigriventris is clothed in ne, erect setae with reddish reections on the head and legs. The elytra are and has a reddish head, pronotum, and legs, and covered with prominently punctate striae. The pronotum is bluish-green elytra. In the male, the length of the narrower than the base of the elytra, which are quadrate. rostrum is shorter than the length of the head and the The head is narrow and the rostrum is almost twice as long antennae are inserted close to the apex. In the female, as the head. The antennae are not elbowed and emerge the rostrum is longer than the head, and the antennae near the apex of the rostrum; the last three antennomeres are inserted near the middle. The head and pronotum are almost twice as wide as the preceding ones. are narrower than the base of the elytra. POLYPHAGA POLYPHAGA

FAMILY FAMILY Brentidae SUBFAMILY SUBFAMILY Brentinae DISTRIBUTION Palearctic: southern Europe, Algeria, Morocco, Israel, Syria, Iran, Russia DISTRIBUTION Nearctic and Neotropical: USA (Florida) to Paraguay MACROHABITAT Temperate humid forests containing predominantly oaks (Quercus spp.) MACROHABITAT Tropical and subtropical MICROHABITAT Nests of Camponotus, Lasius, Crematogaster, Pheidole, Tapinoma, MICROHABITAT Decaying wood; under bark and Myrmica ants FEEDING HABITS Adults feed on sap or visit owers for nectar; larvae bore into FEEDING HABITS Uncertain dead wood and possibly feed on sap or fungal mycelia NOTE Myrmecophilous; has an unusual head morphology NOTE The species exhibits sexual dimorphism and is the largest weevil in North America

ADULT LENGTH ADULT MALE LENGTH /– / in /–1 / in (9–18 mm) (9–50 mm)

AMORPHOCEPHALA CORONATA ANCHORAGO ADULT FEMALE LENGTH /–1/ in AMORPHOCEPHALA (8–27 mm) (LINNAEUS, 1758) 598 CORONATA 599 (GERMAR, 1817)

This species is a facultative ant associate, usually of Most brentid species exhibit sexual dimorphism and Brentus Camponotus but tolerated, after minor hostility, by other anchorago is no exception: Some individuals may be up to ve ant groups. When introduced to a Camponotus colony, these times larger than the smallest. Both sexes engage in combat, brentids face aggression by the workers until the ants discover and those with longer bodies and rostra, which are used as and begin to lick glandular secretions from pubescent areas on weapons, are more successful in securing a mate. There is an the beetle’s head. Apparently, Amorphocephala coronata overall preference by both sexes for larger mates, thus skewing exhibits a pseudoaltruistic behavior, whereby it regurgitates to populations toward larger-bodied individuals. Females chew the colony part of the food received from host worker ants. holes into the decaying wood of primarily Gumbo-limbo Ants have been observed tending to these brentids and actively () trees to oviposit. Adults can be readily trying to retain them in their nest. The species is gregarious, found in large numbers under the bark of dead logs. with many individuals found together. RELATED SPECIES RELATED SPECIES Brentus and Cephalobarus are currently classified in the This genus belongs in the tribe Eremoxenini (which is Neotropical tribe Brentini. Thirty-seven species are included sometimes treated as the subtribe Eremoxenina within the in Brentus. Of these, B. cylindrus has been reported from Brentini), an almost exclusively myrmecophilous group. Polynesia (Marquesas, Tahiti), where it is possible that it was Other genera in this group include Cobalocephalus, introduced. The rst brentid species described by Linnaeus in Eremoxenus, and Symmorphocerus. Amorphocephala contains 1758 were B. anchorago and B. dispar, originally 20 species from the Palearctic and Afrotropical regions. The under the genus Curculio. species can be separated mainly by characteristics of the head, prothorax and antennae.

Amorphocephala coronata is shiny reddish-brown Brentus anchorago is a greatly elongate black beetle with a narrowly elongate body. Its most weevil with reddish-orange vittae on the elytra. outstanding feature is its large, complex head, It is one of the largest weevil species in North with the hind rostrum (just below the frons) deeply America. Males are strikingly longer, with the concave and bearing brushes of sti setae. The head long, slender prothorax narrowing in the middle is sexually dimorphic, the males having a more and the elongate, narrow rostrum about as long robust front rostrum with large sickle-shaped as the prothorax. Females have a tear-shaped mandibles, and the females having elongate, prothorax, broadening basally, and a rostrum cylindrical mandibles. Actual size Actual size that is about half the size of the prothorax.