A Treatise on the Western Hemisphere Caraboidea (Coleoptera): Their Classification, Distributions, and Ways of Life: Trachypachidae, Carabidae - Nebriiformes ... Faunistica) (Pensoft Series Faunistica)

A Treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification,

Pensoft Series Faunistica 66

Terry L. Erwin

Cover: Cicindis horni Bruch

Volume I. Trachypachidae, Carabidae – Nebriiformes 1

Cicindis horni Bruch. Image credit: Valeria Aschero, CRICyT, Mendoza, Argentina.

distributions, and ways of life

synthesis of the classification, distributions, and ways of life of the ground-beetles (Coleoptera: Caraboidea) dwelling in the Western Hemisphere. Under each taxon account, the historical nomenclature, distribution at the country and/or state/province level, and way of life, including Macrohabitat, Microhabitat, Dispersal abilities, Seasonal occurrence, and Behavior are elaborated. Selected references for each taxon are given in these categories and an extensive, nearly complete bibliography of Caraboidea references for the Western Hemisphere is presented. The sections on way of life include the following: Macrohabitat summary, altitudinal distribution in meters, as well as a general altitudinal classification, habitat/microhabitat details, seasonality, and under Behavior notes on diel activity, adult/larval hibernation, prey items, immatures, over wintering, and other notes on behavior are presented. Details of dispersal power, or the capability of dispersal, i.e. wing condition, flight data, walking, running, burrowing and climbing records, as far as is known is included. Species introduced are noted with their probable area of origin. This work, planned as a seven volume companion to Larochelle & LariviØre (2003), will be an indispensable tool for anybody interested in ground-beetles per se and their natural history, as well as their role in the environment and importance to agriculture, conservation, and forestry. National, regional, and local libraries will find this set of tools well used due to the incredible amount of information that is summarized at these geographical levels. This work follows the classification of Erwin (1984), and the nomenclature of Lorenz (2005), as updated on a six month cycle by Erwin (e-data base, NMNH, Smithsonian Institution).

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A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA (COLEOPTERA) THEIR CLASSIFICATION, DISTRIBUTIONS, AND WAYS OF LIFE VOLUME I (TRACHYPACHIDAE, CARABIDAE – NEBRIIFORMES 1)

Terry L. Erwin

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A Treatise on the Western Hemisphere Caraboidea (Coleoptera) Their classification, distributions, and ways of life Volume I (Trachypachidae, Carabidae – Nebriiformes 1)

TERRY L. ERWIN

SOFIA–MOSCOW 2007

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A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA. VOLUME I

A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA (COLEOPTERA) THEIR CLASSIFICATION, DISTRIBUTIONS, AND WAYS OF LIFE VOLUME I (TRACHYPACHIDAE, CARABIDAE – NEBRIIFORMES 1) Terry L. Erwin

First published 2007 ISBN 978-954-642-298-9 Pensoft Series Faunistica No 66 ISSN 1312-0174

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner.

Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359-2-870-42-82 [email protected] www.pensoft.net

Printed in Bulgaria, April 2007

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Dedicated To Professor George Eugene Ball University of Alberta Edmonton, Canada George likes books and little brooks those hard-to-reach places in which carabid beetles dwell; landscapes that make the muscles twitch n’ swell that’s when George Ball hears yon the Beckon Call taking the student – he mentors! shapes! the next generation’s itch to add yet another species to the Tell

GEORGE E. BALL PICO DE NEBLINA, VENEZUELA, 1984 SMITHSONIAN EXPEDITION

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Contents Foreward

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Introduction Methods

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Volume supplements and Database Bibliography Images

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Acknowledgements

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Arrangement of Western Hemisphere taxa: Caraboidea Species accounts

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Family TRACHYPACHIDAE Tribe Systolosomatini, Systolosoma Tribe Trachypachini, Trachypachus

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Family CARABIDAE Tribe Pelophilini, Pelophila 20 Tribe Nebriini, Leistus, Nebria, Nippononebria 22 Tribe Notiokasiini, Notiokasis 51 Tribe Opisthiini, Opisthius 52 Tribe Notiophilini, Notiophilus 53 Tribe Cicindini, Cicindis 61 Tribe Omophronini, Omophron 63 Tribe Loricerini, Loricera 69 Tribe Carabini, Carabus, Calosoma, Callisthenes, Calopachys Subtribe Ceroglossina, Ceroglossus 118 Tribe Cychrini, Cychrus, Scaphinotus, Sphaeroderus 137 Tribe Collyridini, Ctenostoma 171

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General Bibliography for Caraboidea of the Western Hemisphere Index

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Plates

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Foreward Going on nearly 4 decades of admiration for my dear friend and still mentor, to Professor George E. Ball, I dedicate these volumes, hard bound with carabid images and carabid-dwelling landscapes, on the occasion of his 80th birthday (September 28, 2006); to him and to his devoted decades of gathering carabid beetles and data about them, mentoring, apprenticing, training, and teaching young carabid workers and dozens of other-taxa students, building important reference collections, and sharing his time with anyone needing a carabid identification, their distributions, or general life philosophy. Without him, and all those Carabidologists and generalist beetle collectors he trained or influenced, much of what is included in these volumes would not be available to the world of beetle enthusiasts and natural historians, indeed, to the world in general. TERRY L. ERWIN National Museum of Natural History Smithsonian Institution Washington, DC

INTRODUCTION

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Introduction Inspired by the book, A Natural History of the Ground Beetles (Coleoptera: Carabidae) of America North of Mexico (Larochelle, A. & Larivière, M.C. 2003), its utility and elegance on the shelf, and knowing that Prof. George E. Ball prefers books to electronic media, and at the urging of my dear wife Grace who recognized the importance of the Larochelle & Larivière contribution, I decided to migrate the information in my electronic database of Western Hemisphere Caraboidea to the printed page. There are at present 9,242 species and subspecies currently recorded from the Western Hemisphere, therefore this Treatise will necessarily appear in six or seven volumes, with subsequent supplemental updates. The electronic version is updated on a six month basis. These volumes will add to and update the species of North America documented by Larochelle & Larivière (2003) with new information garnered from data in the large collections at NMNH, CAS, CMNH, UASM, and others, as well as my personal field notes from 39 years of neotropical field work, thus extending Larochelle & Larivière’s important contribution south of the U.S. - Méxican border by treating species of northern México, the Neotropics and Neaustral Regions. Together, my volumes and that of Larochelle & Larivière can be considered as a set of handbooks on the ‘ways of life’ of Western Hemisphere carabid beetles. Larochelle & Larivière included information that I don’t cover, for example laboratory feeding and trapping methods of the North American species, while I include information they didn’t include, for example, historical nomenclature, distributions, altitudinal information, and data from specimens in major collections not heretofore published, as well as color images of adults and landscapes/habitats. We both also contribute complementary bibiliographies. Nearly all carabid species need much more study before we can truly know their role in the complexity of earth’s environment. The species’ “way of life snapshots” provided by our combined contributions should be regarded as a starting point for further discoveries, many of which will be exciting and worth telling again and again sometime, somewhere, over a camp fire. The “Tell” (Film: Mad Max Beyond Thunderdome, 1985) began with us in the early evolution of humankind as oral history. Oral history has now evolved into e-sharing of data, e-stories, e-ideas, e-images and a multitude of other types of modern communication. This is the mode by which future generations will ‘tell,’ however, the written page stored amongst dispersed libraries remains, at least today, as the only reliably permanent continuum of human knowledge.

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Methods For over three decades, data on Caraboidea of the Western Hemisphere have been stored in various data management systems in my Tropical Forest Canopy Laboratory at the National Museum of Natural History, Smithsonian Institution. In the early 1990’s, these data were migrated to FileMakerPro and launched on the Department of Entomology website. Updates were made to the website, as reprints were received from colleagues, and at each new issue of the Zoological Record (Coleoptera). Plans are now being made to move this carabid database into SEQUEL, another database management system that is more secure on today’s internet, in anticipation of the Smithsonian’s planned Encyclopedia of Life Project. In addition, a version of the information from each of the six-seven volumes in this series, as they are finished, will be placed on the Tree of Life website in cooperation with D.R. Maddison, (Maddison D.R. (2001) http://tolweb.org). The format for the following species accounts have been standardized, as much as possible. Organization of higher taxa follows that of Erwin (1985) with some modifications based on recent publications. Within each Tribe, genera are arranged alphabetically, and within them, species and subspecies are also arranged alphabetically. General information is presented for the categories Tribe and Genus, including: Number of worldwide genera: Number of Western Hemisphere genera: Taxonomy: Notes on its stability, or lack thereof, and lineages needing revisionary studies. Adelphotaxon: Geographic status: Biogeographic realms. I use standard terminology with the addition of Neaustral (sensu Kavanaugh, pers. comm.) to delineate the Patagonian region. Derivatives of this are Panaustral and Afroaustral. References: Only ones that I used are listed, it is not exhaustive. In addition to the above, I also provide for the Genus: Type species: Distribution: Countries, or realms if widespread, with number of species in each. Habitat: An overview for the genus. Common name (see below) Plates references (these plates are found at the back of the book) Synonym list

METHODS

Table 1. Abbreviations of states and provinces use in the species accounts. AB AG AI AK AL AR AZ BC BJ CA CH CL CM CO CP CS CT DC DE DF DU FL GA GJ GO HD IA ID IL IN JA KS KY LA LB MA MB MD ME MH MI ML MN MO MS MT MX NA

Alberta Aguascalientes Aleutian Islands Alaska Alabama Arkansas Arizona British Columbia Baja California California Chihuahua Coahuila Colima Colorado Campeche Chiapas Connecticut District of Columbia Delaware Distrito Federal Duragngo Florida Georgia Guanajuato Guerrero Hidalgoa Iowa Idaho Illinois Indiana Jalisco Kansas Kentucky Louisiana Labrador Massachusetts Manitoba Maryland Maine Michoacan Michigan Morelos Minnesota Missouri Mississippi Montana México Nayarit

NE NC NB NF NH NJ NK NL NM NS NT NV NY OA OH OK ON OR PA PE PM PQ PU QR QT RI SK SC SD SI SL SO TA TL TM TN TX UT VA VC VT WA WS WV WY YT YC ZA

Nebraska North Carolina New Brunswick Newfoundland New Hampshire New Jersey North Dakota Neuvo Leon New Mexico Nova Scotia Northwest Territories Nevada New York Oaxaca Ohio Oklahoma Ontario Oregon Pennsylvania Prince Edward Island St. Pierre and Miquelon Quebec Puebla Quintana Roo Queretaro Rhode Island Saskatchewan South Carolina South Dakota Sinaloa San Luis Potosi Sonora Tabasco Tlaxcala Tamaulipas Tennessee Texas Utah Virginia Veracruz Vermont Washington Wisconsin West Virginia Wyoming Yukon Territory Yucatan Zacatecas

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An extended focus image or digital image of an exemplar adult of some species in each genus is included and where possible, an exemplar image is provided for each genus of a living beetle and an image of the typical landscape/habitat inhabited by members of the genus. Within each species account, information is presented in the following format: Genus (Subgenus) species name/ author/ date Common name: These names are derived for the most part from the Latin epithet, patronyms, or geographic references. In some cases, I have used the author of the species name in place of the foregoing. Common names are essential for public access to information and particularly in the realm of conservation practices. This is not a new practice; Brullé (1836,1837), for example, provided a common name in French preceding each of his new species and provided common names for those already described by his contemporaries and predacessors. Synonym list: Here I give the original synonymical names arranged in chronological order following the currently accepted name as it was originally proposed. The name is followed by author and date:page citation. Distribution: Here, information about Realms, introductions, countries, and in some cases, States (USA and México) or Provinces (Canada and Argentina) is provided. See Table 1. Way of Life: MACROHABITAT: Here I give a general statement about altitudinal zones, actual meters in altitude, followed by notes on vegetation cover. My protocol is as follows: Lowlands, 0 to 500 meters; midlands, 501 to 1500 meters; uplands, 1501 to 2200 meters; mountains, 2201 to >> n meters. The altitudinal data come directly from specimen labels, or from Google look-ups of specimen localities, if data were not found in publications. Label data given in “feet” were converted to metric. MICROHABITAT: Under this heading is provided as much detailed information as is known from the literature, field notes, or specimen labels regarding strata (ground dwelling, canopy dwelling, etc.), soil types and general moisture content, near or far from water, snow, or other physical feature of the environment wherein members of the species are found. DISPERSAL ABILITIES: Wing length reference; flying, running, burrowing, and climbing abilities. SEASONAL OCCURRENCE: Months in which specimens have been captured. BEHAVIOR: All recorded observations on activities of adults and larvae that I was able to find in the literature and on specimen labels. References: Here I have included pertinent published articles, field notes, specimen label information (i.e. which museum has specimen data referred too), and personal communications from other Carabidologists.

METHODS

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In the following species accounts, a hyphen (-) is used for its normal purpose; a dash (–) is used to indicate a continuum between entries (e.g. between measures, altitudinal ranges, or across months), and for separation between Country and its States/Provinces in the distribution lists. The following codens are used in the species accounts in this volume: CAS, California Academy of Sciences, San Francisco, CA; David H. Kavanaugh, Curator CMNH, Carnegie Museum of Natural History, Pittsburg, PA, Robert Davidson, Curator IADIZA, Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza, Argentina, Sergio Roig-Juñent, Director and Curator NMNH, National Museum of Natural History, Smithsonian Institution, Terry L. Erwin, Curator UASM, University of Alberta, Strickland Museum, Edmonton, Alberta, Canada, George E. Ball and Danny Shpeley, Curators

Volume supplements and Database Upon using this volume and those to follow, readers are encouraged to send notice of any errors, omissions, bibliographic additions, or supply new data and/or images, so that these may be included both in the volume supplementals and on the website database. Full credit will be given to contributors. Please send such to the author at: [email protected].

Bibliography The bibliography attending this first volume is more or less complete for the taxonomy of all carabid species described for the Western Hemisphere plus many articles on way of life. With each ensuing volume, a supplemental bibliography will be added to fill in the gaps and to add newly published papers, as well as more natural history articles.

Images Dorsal aspect mounted beetle images used herein were either made with a Leica M420 microscope with a JVC Digital Camera KY-F7OB attached and processed with GT Vision™ software (refered to in the captions as “extended focus image,” or for larger

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adults, imaged with a Microptics ™ Digital Lab XLT (refered to in the captions as “digital image”.) Leg and antennae orientation differs among habitus images because all were reoriented after removal from pins or points and pin holes rendered in Photoshop™. Background color varies depending on the colors of the specimen being imaged. Figure captions of montage and Microptics imaged specimens include an ADP or other number which is an unique label identification number for the specimen that was illustrated. This number links the specimen and associated illustration and/or images to additional information in electronic databases at the NMNH. Images provided by credited photographers were taken with a variety of different methods; “live” notes in the captions can refer to a specimen actually alive, or to a very freshly dead specimen posed for the image. Plate captions include a measure indicating the ABL (apparent body length, from Ball, 1982) of the specimen montaged or digitally imaged with the Microptics setup. Images of live beetles do not have a scale. These captions also include locality information for individuals imaged, as well as a referenced exemplar landscape/habitat typical for members of the genus. Image credits are provided in each caption. All extended focus images and Microptics images were produced and rendered as necessary in Photoshop by Warren E. Steiner Jr., with the exception of Notiokasis (see caption).

ACKNOWLEDGEMENTS

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Acknowledgements First and foremost, heartfelt and gracious thanks go to George E. Ball for my apprenticeship into the world of carabid beetles and his continuing role as my mentor and amigo; and, to his colleague and fellow Carabidologist, Danny Shpeley, whose dedicated work on the Méxican collection (UASM) contributed significantly to this Treatise; and, to Andre L. Larochelle and Marie Claude Larivière for stimulating this effort and allowing their data to be incorporated into the website behind this Treatise, as well as the liberal use of their prose on North American carabids in this book. To Grace P. Servat (Erwin) who saw the book, A natural history of the ground-beetles (Coleoptera: Carabidae) of America north of Mexico by Larochelle and Larivière, and showed me the way toward getting my electronic data base out of “e” and into this published Treatise; a gigantic hug for her. Carl H. Lindroth led the way first, grandly and systematically, for all of us with his treatment of the ground beetles of Newfound, and later in his Magnum Opus of Canada and Alaska in which he presented a snapshot of the ‘way of life’ for many of his included one-thousand plus species. Larochelle and Larivière captured and enhanced much of this for their book and dedicated it to Carl; I captured their enhancement and built on it. Our debt to Carl is enormous. Special thanks also go to George Venable who got me into using FileMakerPro and the resulting data base on which this Treatise is built; to Warren E. Steiner who provided the extended focus images and Microptics images for this volume plus many trips to the library, and to Stephen McJonathan of GT Vision for the loan of equipment and software necessary to make the extended focus images; to Roy Anderson, Allan Ashworth, Christopher J. Earle, Henri Goulet, David Kavanaugh, David Maddison, and Grace Servat who provided other images which are credited appropriately in the figure captions; to many colleagues who sent their publications to the Smithsonian Carabid Library; and to David H. Kavanaugh, Roberta Brett, and Allen Ashworth for their valuable contributions to the Nebriini, Omophronini, and Trachypachidae sections, respectively. And, a hearty thanks to Sergio Roig-Juñent who personally checked the identifications of Ceroglossus species in the NMNH and to Soledad Sallenave who did some library work for the Patagonian references.

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Arrangement of Western Hemisphere taxa1 Caraboidea Volume 1 Family TRACHYPACHIDAE Tribe Systolosomatini, Systolosoma Tribe Trachypachini, Trachypachus Family CARABIDAE Division NEBRIIFORMES Subfamily CARABINAE Supertribe NEBRIITAE Tribe Pelophilini, Pelophila Tribe Nebriini, Leistus, Nebria, Nippononebria Tribe Notiokasiini, Notiokasis Tribe Opisthiini, Opisthius Tribe Notiophilini, Notiophilus Supertribe CICINDITAE Tribe Cicindini, Cicindis Supertribe OMOPHRONITAE Tribe Omophronini, Omophron Supertribe LORICERITAE Tribe Loricerini, Loricera Supertribe CARABITAE Tribe Carabini, Carabus, Calosoma, Callisthenes, Calopachys, Ceroglossus [Tribe Pamborini] Tribe Cychrini, Cychrus, Scaphinotus, Sphaeroderus Supertribe CICINDELITAE Tribe Collyridini, Ctenostoma

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Name in brackets is strictly an Australian taxon included here only for completeness of Tribal list.

SPECIES ACCOUNTS / TRACHYPACHIDAE

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SPECIES ACCOUNTS TRACHYPACHIDAE FALSE GROUND BEETLES Systolosomatini Erwin 1985 NEAUSTRAL FALSE GROUND BEETLES Systolosomatini Erwin, 1985:467

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Trachypachini Thomson 1857. Geographic status: Neaustral. References: Erwin (1985, 2001) Systolosoma Solier 1849 NEAUSTRAL FALSE GROUND BEETLES (Plate 1) Systolosoma Solier, 1849:241 Notioxenus Motschulsky, 1857:111

Number of species: Two. Taxonomy: Stable. Adelphotaxon: Trachypachus Motschulsky 1844. Type species: Systolosoma breve Solier, 1849. Distribution: Native, New World: ARGENTINA, Chile. (Two species) Habitat: Neaustral Beech and conifer forests, thickets, and coppice. References: Ashworth (pers. comm.), Erwin (2001), Nègre (1973) Systolosoma breve Solier 1849 SHORT NEAUSTRAL FALSE GROUND BEETLE Systolosoma breve Solier, 1849:242 Notioxenus bilunulatus Motschulsky, 1857:111

Distribution: Native, New World. ARGENTINA – NEUQUÉN, RÍO NEGRO; CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 40 – 1350 meters altitude, in Valdivian rainforest with Nothofagus dombeyi (Mirb.) Blume and Araucaria araucana (Molina) K. Koch, CoigueManiu Forest, and Nire/Steppe. MICROHABITAT: On sandy alluvium and open sand, as well as in forest leaf and wood litter. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – March, and October – December. BEHAVIOR: Adults take cover in leaf litter and are attracted to dung-baited and squid-baited pitfall traps. They fly low to the ground and are often caught in window traps.

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References: Ashworth (pers. comm.), Erwin (2001), Data from CAS, NMNH collections Systolosoma lateritium Nègre 1973 TILED NEAUSTRAL FALSE GROUND BEETLE Systolosoma lateritius Nègre, 1973:291

Distribution: Native, New World. ARGENTINA – RÍO NEGRO: CHILE. Way of Life: MACROHABITAT: Midlands, 530 – 1350 meters altitude, Valdivian temperate rain forests, and in Coigue-Maniu forest, Coigue-Lenga forest, Lenga Beech forest, (Nothofagus dombeyi (Mirb.) Blume-N. pumilio (Poepp & Endl.) Krasser forests, N. pumilio forests (w/ Chusquea), and at tree line. MICROHABITAT: On open ground, as well as in forest leaf and wood litter. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January, March, and October – December. BEHAVIOR: Adults take cover in leaf litter and are attracted to dung-baited pitfall traps. References: Ashworth (pers. comm.), Erwin (2001), Nègre (1973), Data from CAS, NMNH collections Trachypachini Thomson 1857 TEMPORAL FALSE GROUND BEETLES Trachypachini Thomson, 1857:5

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Systolosomatini Erwin 1985. Geographic status: Holarctic. Distribution: Canada, northwestern USA, northern Europe. References: Erwin (2001), Lindroth (1961) Trachypachus Motschulsky 1844 ROUGH TEMPORAL FALSE GROUND BEETLES (Plate 2) Trachypachus Motschulsky, 1844:86 Trachypachys Gemminger & Harold, 1868:46

Number of species: Four. Taxonomy: Stable. Adelphotaxon: Systolosoma Solier 1849. Type species: Blethisa Zetterstedtii Gyllenhal, 1827. Distribution: Native, New World (three species), northern Europe (one species). Habitat: Sandy or dusty ground beneath deciduous or conifer forests, or on open faced embankments of exfoliating clay. References: Erwin (2001), Lindroth (1961), Maddison (2001)

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Trachypachus gibbsii LeConte 1861 GIBBS’ TEMPORAL FALSE GROUND BEETLE Trachypachus Gibbsii LeConte 1861:339 Trachypachus californicus Motschulsky, 1864:194 Trachypachus alticola Casey, 1920:144

Distribution: Native, New World. CANADA – BC; USA – CA, ID, MT, OR, WA. Way of Life: M ACROHABITAT : Midlands to mountains, 762 – 3505 meters altitude. MICROHABITAT: Adults of this species are stenotopic, restricted to a special habitat where it often is abundant: on the banks of large rivers and small forested creeks, often some distance from the water. Also, they occur under poplars (Populus balsamifera L.) where the ground, half shaded, consists of fine dusty dry sand covered with dry leaves, no herb layer present, and in pine/fir forests in similar soil conditions. DISPERSAL ABILITIES: Macropterous, and recorded in flight. SEASONAL OCCURRENCE: Adults have been found in March, May – September, and November. BEHAVIOR: Adults run in the sunshine on open bare spots; they have been extracted from Noble Fir (Abies procera Rehd.) duff using Berlese funnels. References: Erwin (2001), Lindroth (1961), Data from CAS, NMNH collections Trachypachus holmbergi von Mannerheim 1853 HOLMBERG’S TEMPORAL FALSE GROUND BEETLE Trachypachus Holmbergi von Mannerheim, 1853:119 Trachypachus inermis Motschulsky, 1864:194 Trachypachus oregonus Casey, 1920:145 Trachypachus specularis Casey, 1920:146

Distribution: Native, New World. CANADA – AB, BC, NT, SK, YT; USA – AK, CA, CO, ID, MT, NV, OR, WA. Way of Life: MACROHABITAT: Lowlands to mountains, 5 – 2652 meters altitude. MICROHABITAT: Often among dead leaves. According to Maddison (2001), adults of Trachypachus holmbergi are “found on loose loam or conifer duff. For example, specimens were common in one regularlytilled garden at the University of Alberta. They were found in a similar, but more natural habitat, at the base of a slope on Cape Perpetua, Oregon”. Adults of this species are xerophilus, occurring independent of water and are found on open or slightly shaded ground, usually moraine, with sparse vegetation. DISPERSAL ABILITIES: Macropterous, and these no doubt functional. SEASONAL OCCURRENCE: Adults have been found in April – August, October, and December. BEHAVIOR: Adults run in the sunshine, and dive into the substrate to escape capture. Adults hibernate. References: Erwin (2001), Lindroth (1961), Maddison (2001), Data from CAS, NMNH collections Trachypachus slevini Van Dyke 1925 SLEVIN’S TEMPORAL FALSE GROUND BEETLE Trachypachus slevini Van Dyke, 1925:111

Distribution: Native, New World. USA – OR, WA. Way of Life: MACROHABITAT: Lowlands to midlands, sea level–1045 meters altitude. MICROHABITAT: “At Moolack Beach along the coast of Oregon, Trachypachus slevini can be found

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running on the cliff above the sandy beach of the Pacific Ocean. The beetles run in the talus slope, consisting of small particles, at the base of the cliff of exfoliating clay. The beetles run along for a short way in the sunshine, then dive into the pile of debris; they also are found higher up on the cliff on the hard clay.” Maddison (2001). DISPERSAL ABILITIES: Macropterous, and these no doubt functional. SEASONAL OCCURRENCE: Adults have been found in July – October. BEHAVIOR: Adults take cover under debris on the ground. References: Erwin (2001), Maddison (2001), Data from NMNH collection

CARABIDAE GROUND BEETLES NEBRIIFORMES, PART 1 CARABINAE NEBRIITAE

Pelophilini Kavanaugh 1996 MUD-LOVING BEETLES Pelophilini Kavanaugh, 1996:35

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: (Notioksini + Opisthiini + Nebriini) Geographic status: Holarctic. References: Erwin (2001), Lindroth (1961), Kavanaugh (1996, 1999) Pelophila Dejean 1826 MUD-LOVING BEETLES (Plate 3) Pelophila Dejean, 1826:262

Number of species: Two. Taxonomy: Stable. Adelphotaxon: See Pelophilini, above. Type species: Carabus borealis Paykull, 1790. Distribution: Native, New World (one species), Palearctic + Boreal North America (one species). Habitat: Forest swamps, above tree line, and on the tundra, always near water in open situations on soft ground with a rich content of organic matter. References: Erwin (2001), Lindroth (1961), Kavanaugh (1996, 1999) Pelophila borealis (Paykull) 1790 BOREAL MUD-LOVING BEETLE Carabus borealis Paykull, 1790:61 Pelophila elongata, von Mannerheim, 1823:38 Pelophila Eschscholtzii, von Mannerheim, 1823:40

SPECIES ACCOUNTS / CARABIDAE

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Pelophila Gebleri, von Mannerheim, 1823:38 Pelophila marginata, von Mannerheim, 1823:38 Pelophila Arctica, Dejean, 1826:265 Pelophila Dejeanii, Dejean, 1826:265 Pelophila laevigata, Motschulsky, 1844:92 Pelophila californica, Motschulsky, 1844:93 Pelophila angusticollis, Motschulsky, 1844:98 Pelophila ochotica, R.F. Sahlberg, 1844:17 Pelophila costata, Ménétriés, 1851:49 Pelophila Ulkei, G. Horn, 1870:105 Pelophila Shermani, Casey, 1913:45

Distribution: PALEARCTIC; CANADA – AB, BC, LB, MB, NF, NT, NU, ON, PQ, SK, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands to midlands, subalpine and alpine zones, 10 – 590 meters altitude, at the borders of slow rivers, lakes, eutrophic ponds, pools, marshes, and swamps. MICROHABITAT: Adults are markedly hygrophilous, always near water in open situations, occurring also above tree line and on the true tundra. Found on soft ground with rich content of organic matter, usually in dense vegetation (e.g. Juncus balticus sitchensis, Carex aquatilis, Eriophorum angustifolium, Arctophila fulva & Hippuris, sometimes in moss carpets of Campylium stellatum, & co). DISPERSAL ABILITIES: Macropterous. Flight recorded in Europe. Adults and larvae are fast runners; adults are excellent swimmers. SEASONAL OCCURRENCE: Adults are active in May – September (November record: LB). Gravid females found in late June and early July (PQ). Eggs are laid in the ground. Teneral adults emerge in early July (PQ). BEHAVIOR: Nocturnal, predatory in both adult and larval stages feeding on worms, insect larvae, and carrion; adults take cover during the day under vegetal debris, stones, and in moss carpets. They are able to withstand freezing temperatures (as low as -5°C). They have been found over wintering in cavities in the ground in Europe. Larval habitat: under stones. Adults hibernate. References: Erwin (2001), Kavanaugh, (in litt.), Larochelle and Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Pelophila rudis (LeConte) 1863 WILD MUD-LOVING BEETLE Nebria rudis LeConte, 1863:3

Distribution: Native, New World. CANADA – AB, BC, MB, NF, NT, ON, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands, 150 – 620 meters altitude, in temporary swamps with Alnus rugosa (Du Roi) Sprengel, Salices, & co. MICROHABITAT: Adults are found in dense, wet carpets of Drepanocladus aduncus, with Naumburgia thyrsiflora, Triglochin palustre, Eleocharis palustris, Carices, Juncus, & co. at the margins of eutrophic marshes; marshy borders of slow streams, in swamps, and in lakeside forests near sandy lake shores. They are local. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Adults are fast runners. SEASONAL OCCURRENCE: Adults are found in May – July, September – November, teneral adults in September (AB). Found in stumps and fallen trunks in shady spots in May in Newfoundland. BEHAVIOR: Nocturnal, predatory in both adult and larval stages. Adults hibernate. References: Erwin (2001), Kavanaugh, (in litt.), Larochelle and Larivière (2003), Lindroth (1961), Data from NMNH collection

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Nebriini Castelnau de Laporte 1834 GAZELLE BEETLES Nebriini Castelnau de Laporte, 1834:90

Number of worldwide genera: Three. Number of Western Hemisphere genera: Three. Taxonomy: In flux; phylogenetic analysis of the worldwide fauna may change the number of genera recognized. Adelphotaxon: (Notiokasini + Notiophilini + Opisthiini) Geographic status: Holarctic. References: Erwin (2001), Lindroth (1961). Notes: Presentation of this tribe was significantly enhanced by published and unpublished information shared by David H. Kavanaugh, CAS, as well as suggestions for many of the common names. Leistus Frölich 1799 BASKET-THROATS (Plate 4) Leistus Frölich, 1799:9 Leistidius K. Daniel, 1903:171 Leistophorus Reitter, 1905:213 Euleistulus Reitter, 1905:224 Acroleistus Reitter, 1905:225 Neoleistus Erwin, 1970:112 [valid subgenus for native Nearctic species]

Number of worldwide species: 181. Number of Western Hemisphere species: Four. Taxonomy: Stable in the Nearctic, many new species yet to be discovered in Asia. There is no synthesis of Palearctic taxa. Adelphotaxon: Nipponnebria Ueno 1955. Type species: Leistus testaceus Frölich, 1799. Distribution: Native, New World (three species), introduced (one species), Palearctic (136 species). Habitat: In lowlands and mountainous areas. Found in open woodlands and near water in open situations on soft ground with a rich content of organic matter, or in moist gravelly ground. References: Erwin (1970, 2001), Kavanaugh (1998), Lindroth (1961), Larochelle & Larivière (2003) Notes: The common name of this genus, Basket-Throats, was suggested by D.H. Kavanaugh (pers. comm.) who will be using this name in upcoming publications. Leistus (Leistus) ferrugineus (Linné) 1758 PALEARCTIC RUSTY BASKET-THROAT Carabus ferrugineus Linné, 1758:415 Carabus spinilabris Panzer, 1797: no. 11 Leistus testaceus Frölich, 1799:8 Leistus kulti Smetana, 1952:234

Distribution: Introduced. PALEARCTIC; CANADA – NF.

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Way of Life: MACROHABITAT: Lowlands, 10 meters altitude, in mixed forest (Abies, Alnus, Prunus). MICROHABITAT: On moist gravelly ground and grassy areas, hedges, and in open woodlands. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – July and August – October. BEHAVIOR: Nocturnal, adults take cover during the day under stones, in leaf litter and in humus. Adult are predaceous on insects. They are likely larval hibernators, as are all other known European species. References: Erwin (1970, 2001), Lindroth (1961), Larochelle & Larivière (2003), Data from UASM collection Leistus (Neoleistus) ferruginosus von Mannerheim 1843 NEARCTIC RUSTY BASKET-THROAT Leistus ferruginosus von Mannerheim, 1843:187 Leistus Ferrugineus Dejean, 1831:569 Leistus nigropiceus Casey, 1913:45

Distribution: Native, New World. CANADA – AB, BC, QC; USA – AK, CA, MT, OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 30 – 1830 meters altitude, in deciduous forest. MICROHABITAT: Adults occur on moderately moist, half-shaded ground, usually near running water in leaf litter under Alnus and Salix. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Adults are fast runners and good climbers. SEASONAL OCCURRENCE: Adults are found in February, April – October; teneral adults in June in BC. BEHAVIOR: Adults climb on the foliage of several Vaccinium species in understory vegetation in lowland “meadow forest” at night. They are likely larval hibernators, as are all known European species. References: Erwin (1970, 2001), Kavanugh (in litt.), Lindroth (1961), Larochelle & Larivière (2003), Data from UASM collection Leistus (Neoleistus) longipennis Casey 1920 LONG-WINGED BASKET-THROAT Leistus longipennis Casey, 1920:148

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands and coastal foothills, 15 – 140 meters altitude, in coniferous redwood forests (Sequoia sempervirens (D. Don) Endl. Taxodiaceae). MICROHABITAT: Found at night climbing thimbleberry (Rubus parviflorus Nutt. Rosaceae) at margins of forests, or along stream margins. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Adults are fast runners and good climbers on low vegetation; found up to meter above soil surface. SEASONAL OCCURRENCE: Adults have been found in March, and May – August; teneral adults in June – July. BEHAVIOR: They are likely larval hibernators, as are all known European species. References: Erwin (1970, 2001), Kavanugh (in litt.), Larochelle & Larivière (2003), Data from CAS, NMNH collections

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A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA. VOLUME I

Leistus (Neoleistus) madmeridianus Erwin 1970 MAD RIVER BASKET-THROAT Leistus madmeridianus Erwin, 1970:117

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands and coastal foothills, 30 – 90 meters altitude, in coniferous redwood forests (Sequoia sempervirens (D. Don) Endl. Taxodiaceae). MICROHABITAT: Likely the same as L. longipennis Casey, but south of the Mad River, for example in leaf litter under streamside shrubs at Jacoby Creek, near Arcada, CA. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Adults are fast runners and good climbers. SEASONAL OCCURRENCE: Adults have been found in March – April, June – July, November – December; teneral adults in June – July. BEHAVIOR: They are likely larval hibernators, as are all known European species. References: Erwin (2001), Erwin (1970), Kavanugh (in litt.), Lindroth (1961), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Nebria Latreille 1802 GAZELLE BEETLES (Plate 5) Nebria Latreille, 1802:221 Boreonebria Jeannel, 1937:2 [valid subgenus for native Nearctic and some Palearctic species] Reductonebria Shilenkov, 1975:87 [valid subgenus for native Nearctic and some Palearctic species] Catonebria Shilenkov, 1975:88 [valid subgenus for native Nearctic and some Palearctic species] [See Lorenz (2005) for additional Old World taxa]

Number of worldwide species: 383 (+ 158 subspecies). Number of Western Hemisphere species: 53 (+ 48 subspecies). Taxonomy: Stable in the Nearctic, many new species are yet to be discovered in Asia. There is a recent synthesis of World taxa by Ledoux & Roux (2005). Adelphotaxon: Nippononebria + Leistus + Archastes + Eurynebria. Type species: Carabus brevicollis Fabricius, 1792. Distribution: Native, New World (52 species); introduced (one species), Palearctic (more than 335 species). Habitat: According to Lindroth (1961), most species in this genus are markedly hygrophilus, usually confined to stoney, barren margins of running water; some are also found on upper sea beaches, or above treeline in open dry areas, or near snow patches and rills (Kavanaugh, 1992). Adults are carnivorous and nocturnal. Many species are flightless. References: Erwin (2001), Erwin & Ball (1972), Kavanaugh (1971, 1979, 1981, 1984, 1992), Larochelle & Larivière (2003), Lindroth (1961), Lorenz (2005) Nebria (Reductonebria) acuta acuta Lindroth 1961 SHARP-SHOULDERED GAZELLE BEETLE Nebria acuta acuta Lindroth, 1961:71

Distribution: Native, New World. CANADA – BC; USA – AK, CA, OR, WA.

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Way of Life: MACROHABITAT: Lowland to upland forests, subalpine and alpine zones, 10 – 2130 meters altitude. MICROHABITAT: Adults are markedly hygrophilous, usually confined to stony, barren margins of running water, or near snow patches and rills. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in late August. BEHAVIOR: Adults are nocturnal, carnivorous, and hibernation, at least in part, takes place in the adult stage. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Notes: The common name of this species, SHARP-SHOULDERED GAZELLE BEETLE, was suggested by D.H. Kavanaugh (pers. comm.) because Lindroth was referring to the acute humeri found in the adults. Kavanaugh will be using this name in his upcoming publications on this genus. Nebria (Reductonebria) acuta quileute Kavanaugh 1979 QUILEUTE GAZELLE BEETLE Nebria acuta quileute Kavanaugh, 1979:98

Distribution: Native, New World. USA – WA. Way of Life: MACROHABITAT: Midlands to uplands, 560 – 1830 meters altitude. MICROHABITAT: River banks and near springs on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.) Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) acuta sonorae Kavanaugh 1981 SONORA PASS GAZELLE BEETLE Nebria acuta sonorae Kavanaugh, 1981:438

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 2440 – 3320 meters altitude. MICROHABITAT: Streams banks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1981, 1985, in litt.) Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) appalachia Darlington 1931 SOUTHERN APPALACHIAN GAZELLE BEETLE Nebria appalachia Darlington, 1931:153

Distribution: Native, New World. USA – NC, TN. Way of Life: MACROHABITAT: Midlands to uplands, 910 – 2010 meters altitude, in mixed forests. MICROHABITAT: Banks of small forest streams on wet, gravelly ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October; teneral adults in early July. BEHAVIOR:

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Adults are nocturnal, carnivorous. They take cover under stones during the daylight hours. Hibernation, at least in part, takes place in the adult stage. References: Erwin (2001), Barr (1969), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) arkansana arkansana Casey 1913 ARKANSAS RIVER GAZELLE BEETLE Nebria arkansana arkansana Casey, 1913:52

Distribution: Native, New World. USA – CO, NM, UT, WY. Way of Life: MACROHABITAT: Uplands to mountains, subalpine and alpine zones, not above treeline in Canada, 2010 – 3810 meters altitude. MICROHABITAT: Confined to stony-gravelly margins of fast running waters, usually small brooks; DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous. Larvae and adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) arkansana edwardsi Kavanaugh 1979 EDWARDS’ GAZELLE BEETLE Nebria arkansana edwardsi Kavanaugh, 1979:100

Distribution: Native, New World. CANADA – AB, BC, YT; USA – ID, MT, NV, OR, WA, WY. Way of Life: M ACROHABITAT : Midlands to mountains, 720 – 3200 meters altitude. MICROHABITAT: Margins of cool streams on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September; copulating pairs in August. BEHAVIOR: Adults are nocturnal and carnivorous, adults take cover in the day under stones and drift. Larvae and possibly adults hibernate. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) arkansana fragilis Casey 1924 FRAGILE GAZELLE BEETLE Nebria arkansana fragilis Casey, 1924:21 Nebria uinta Kavanaugh, 1979:102 Nebria teewinot Kavanaugh, 1979:103

Distribution: Native, New World. USA – UT, WY. Way of Life: MACROHABITAT: Midlands to mountains, 1490 – 3470 meters altitude. MICROHABITAT: Confined to margins of cool streams on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal and carnivorous, adults take cover in the day under stones and drift. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

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Nebria (Reductonebria) arkansana oowah Kavanaugh 1979 LA SAL MOUNTAINS GAZELLE BEETLE Nebria arkansana oowah Kavanaugh, 1979:102

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Mountains, 2680 – 2900 meters altitude. MICROHABITAT: Confined to margins of cool streams on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Nebria) brevicollis (Fabricius) 1792 SHORT-COLLARED GAZELLE BEETLE Carabus brevicollis Fabricius, 1792:150 Carabus cursor O.F. Müller, 1776:78 Carabus rufipes Goeze, 1777:622 Carabus infida Rossi, 1792:88 Nebria rugimarginata Marsham, 1802:444 Nebria fuscata Bonelli, 1810:64 Nebria lata Newman, 1833:284 Nebria varicornis Newman, 1833:285 Nebria elongata Fischer von Waldheim, 1828:266 Nebria rufomarinata Fischer von Waldheim, 1828:248 Nebria Tamsii Dejean, 1831:573 Nebria barbara Chaudoir, 1843:748 Nebria rufipes Dalla Torre, 1877:24 Nebria cursor Bedel, 1879:17 Nebria sicula Preudhomme de Borre & Ragusa, 1882:181 Nebria obscurata J.R. Sahlberg, 1903:3 Nebria uzokensis Obenberger, 1917:9

Distribution: Palearctic; CANADA – PM, PQ. Introduced, eurytopic; probably not established. Way of Life: MACROHABITAT: Lowlands, sea level – 50 meters altitude in North America. MICROHABITAT: In Europe, this species commonly dwells in partially shaded, hedgerow or woodland habitats such as deciduous forests and parks and often invades adjacent agricultural lands. They prefer moist mull ground in beech forests. It is one of the most abundant species in disturbed or synanthropic situations. Absent only from very saline or very wet situations. Replaced by Nebria salina in some drier habitats, including peatlands and the summit moraine of mountains. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in June in North America. BEHAVIOR: Adults are nocturnal, carnivorous, and autumn breeders. Both adults and larvae hibernate. Adults emerge in the spring and after a short period of activity they enter summer dormancy, aggregating under bark of tree stumps, under fallen trees in the forest, etc. Activity resumes in the fall. References: Erwin (2001), Kavanaugh (in litt.), Larochelle & Larivière (2003), Lindroth (1961)

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Nebria (Catonebria) calva Kavanaugh 1984 MOUNT BALDY GAZELLE BEETLE Nebria calva Kavanaugh, 1984:164

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Mountains, 2710 – 3350 meters altitude, in coniferous forests (Picea, Abies & co.). MICROHABITAT: According to Kavanaugh (in litt.), under stones along cold streams in upper forested and treeline areas. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) carri Kavanaugh 1979 CARR’S GAZELLE BEETLE Nebria carri Kavanaugh, 1979:107

Distribution: Native, New World. USA – ID, MT. Way of Life: M ACROHABITAT : Uplands to mountains, 2070 – 2590 meters altitude. MICROHABITAT: Margins of cool springs on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) catenata Casey 1913 SOUTHERN CHAIN-BACKED GAZELLE BEETLE Nebria catenata Casey, 1913:49

Distribution: Native, New World. USA – CO, UT. Way of Life: M ACROHABITAT : Uplands to mountains, 2000 – 3350 meters altitude. MICROHABITAT: Margins of rivers and cool springs on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) charlottae Lindroth 1961 GRAHAM ISLAND GAZELLE BEETLE Nebria charlottae Lindroth, 1961:67

Distribution: Native, New World. CANADA – QC. Way of Life: MACROHABITAT: Lowlands, 2 – 10 meters altitude. According to Kavanaugh (1992), adults of this species are restricted to north-facing, cobble-type upper sea beaches that are at least partially shaded by the adjacent forest edge. MICROHABITAT: Stenotopic adults have only been found where the cobble is piled at least six inches deep and freshwater seeps through

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the sand/gravel substrate below. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Some beaches on which this species occurs are strewn with driftwood, and adults are found in great numbers running, feeding, and copulating on these fallen trees at night; adults take cover in the day under cobble. References: Erwin (2001), Kavanaugh (1992, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) coloradensis Van Dyke 1943 COLORADO GAZELLE BEETLE Nebria coloradensis Van Dyke, 1943:19

Distribution: Native, New World. USA – CO. Way of Life: MACROHABITAT: Mountains, 2870 – 3350 meters altitude. MICROHABITAT: Found on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Boreonebria) crassicornis crassicornis Van Dyke 1925 THICK-HORNED GAZELLE BEETLE Nebria crassicornis crassicornis Van Dyke, 1925:121

Distribution: Native, New World. CANADA – BC; USA – WA. Way of Life: MACROHABITAT: Midlands to uplands, 1190 – 2190 meters altitude. MICROHABITAT: Not confined to the vicinity of water, but found in the woods, at the margins of bogs and near snow patches on moist or wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Boreonebria) crassicornis intermedia Van Dyke 1949 INTERMEDIATE GAZELLE BEETLE Nebria crassicornis intermedia Van Dyke, 1949:49

Distribution: Native, New World. CANADA – AB, BC; USA – ID, MT, OR, UT, WA, WY Way of Life: MACROHABITAT: Midlands to mountains, alpine zone, 1220 – 3470 meters altitude. MICROHABITAT: Not strictly riparian, but found in the vicinity of water; adults are found in open meadows near snow patches at about the treeline, in moderately dense conifer forests, and near lakes, always on dry ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adults are nocturnal, carnivorous. Hibernation takes place in both the adult and larval stages. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

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Nebria (Reductonebria) danmanni Kavanaugh 1981 DAN MANN’S GAZELLE BEETLE Nebria danmanni Kavanaugh, 1981:437

Distribution: Native, New World. USA – WA. Way of Life: MACROHABITAT: Uplands, 1770 – 1980 meters altitude. MICROHABITAT: Confined to margins of cold rills adjacent to and beneath glaciers on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July and September. BEHAVIOR: At night on top of the glaciers preying on half frozen insects. Adults are nocturnal, carnivorous, taking cover in the day under glaciers in rubble. References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) darlingtoni Kavanaugh 1979 DARLINGTON’S GAZELLE BEETLE Nebria darlingtoni Kavanaugh, 1979:104

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Midlands, 980 – 1340 meters altitude. MICROHABITAT: On river banks and near waterfalls on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) desolata Kavanaugh 1971 DESOLATE GAZELLE BEETLE Nebria desolata Kavanaugh, 1971:41

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Upland desert, 1710 – 2130 meters altitude. MICROHABITAT: At the base of steep reddish sandstone cliffs on cool, wet shaded ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April, and June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under blocks of sandstone. References: Erwin (2001), Kavanaugh (1971, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) diversa LeConte 1863 SEA BEACH GAZELLE BEETLE Nebria diversa LeConte, 1863:2 Nebria livida LeConte, 1859:84 Nebria Townsendi Casey, 1924:19

Distribution: Native, New World. Canada – BC, QC; USA – CA, OR, WA.

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Way of Life: MACROHABITAT: Lowlands, sea level – 5 meters altitude. According to Kavanaugh (1992), this species is restricted to open sandy sea beaches. MICROHABITAT: Adults and larvae are found beneath driftwood and other debris above the highest tide line, especially where sand dunes have developed above the beaches. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – December; teneral adults in May – June, late July. BEHAVIOR: Adults are nocturnal, carnivorous, and semi-gregarious, taking cover in cliff bank crevices some distance from the water. Larvae hide under driftwood. Larvae hibernate. References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) eschscholtzii Ménétriés 1843 ESCHSCHOLTZ’S GAZELLE BEETLE Nebria Eschscholtzii Ménétriés, 1843:55 Nebria tenuipes Casey, 1913:51 Nebria transversa Casey, 1920:152 Nebria formalis Casey, 1920:153 Nebria pallidissima Casey, 1924:19 Nebria pugetana Casey, 1924:19

Distribution: Native, New World. USA – CA, ID, NV, OR, WA Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1650 meters altitude. MICROHABITAT: At edges of small to large streams on silty, sandy, or gravelly substrate. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Adults are nocturnal, carnivorous, and take cover under stones or debris in the day. Adults overwinter or aestivate. References: Erwin (2001), Kavanaugh (1985, in litt.), Lindroth (1961) Nebria (Boreonebria) frigida R.F. Sahlberg 1844 FRIGID GAZELLE BEETLE Nebria frigida R.F. Sahlberg, 1844:11 Nebria viridis G. Horn, 1870:101 Nebria parvula J.R. Sahlberg, 1887:47 Nebria reducta Casey, 1920:150

Distribution: PALEARCTIC; CANADA – BC, NT, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands to midlands, 140 – 1100 meters altitude. MICROHABITAT: Moderately hygrophilus, but not confined to the borders of permanent waters, though adults always occur on rather wet ground. On river banks, they prefer the cover of shrubs and trees, often in the dead leaves. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. They occur up to and slightly above treeline, but this species is not a member of the true tundra fauna. Larvae hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Catonebria) gebleri albimontis Kavanaugh 1984 WHITE MOUNTAINS GAZELLE BEETLE Nebria gebleri albimontis Kavanaugh, 1984:163

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 2320 – 3410 meters altitude. MICROHABITAT: Found on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July and September. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1984, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) gebleri cascadensis Kavanaugh 1979 CASCADE MOUNTAINS GAZELLE BEETLE Nebria gebleri cascadensis Kavanaugh, 1979:105

Distribution: Native, New World. CANADA – BC; USA – OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 300 – 1680 meters altitude. MICROHABITAT: On river banks, at brook margins, and on lake shores on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) gebleri fragariae Kavanaugh 1979 STRAWBERRY MOUNTAINS GAZELLE BEETLE Nebria gebleri fragariae Kavanaugh, 1979:106

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Midlands to uplands, 1490 – 2070 meters altitude. MICROHABITAT: At brook margins, on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May and July. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) gebleri gebleri Dejean 1831 GEBLER’S GAZELLE BEETLE Nebria Gebleri Dejean, 1831:573 Nebria melanaria Hatch, 1949:115

Distribution: Native, New World. CANADA – AB, BC, YT; USA – AK, ID, MT, OR, WA. Way of Life: MACROHABITAT: Lowlands to mountains, 50 – 2590 meters altitude. MICROHABITAT: At stream margins, close to the water on wet, stony, bare ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. Adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Catonebria) gebleri rathvoni LeConte 1853 RATHVON’S GAZELLE BEETLE Nebria Rathvoni LeConte, 1853:400

Distribution: Native, New World. USA – CA, NV. Way of Life: M ACROHABITAT : Midlands to mountains, 910 – 3200 meters altitude. MICROHABITAT: On river banks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) gebleri siskiyouensis Kavanaugh 1979 SISKIYOU GAZELLE BEETLE Nebria gebleri siskiyouensis Kavanaugh, 1979:107

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Midlands to uplands, 700 – 1750 meters altitude. Not recorded from above tree line. MICROHABITAT: Along mountain streams on forested slopes, often quite small brooks, with stony, barren banks; always close to the water. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. Larvae hibernate. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Boreonebria) gouleti Kavanaugh 1979 GOULET’S GAZELLE BEETLE Nebria gouleti Kavanaugh, 1979:94

Distribution: Native, New World. USA – ID, OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 320 – 1800 meters altitude. MICROHABITAT: Moderately hygrophilus and found on banks of rivers, brooks, and near waterfalls on rather wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) gregaria Fischer von Waldheim 1820 GREGARIOUS GAZELLE BEETLE Nebria gregaria Fischer von Waldheim, 1820: pl. 6 Nebria cuneata Casey, 1913:50

Distribution: Native, New World. USA – AK. Way of Life: MACROHABITAT: Lowlands, sea level – 46 meters altitude. MICROHABITAT: Restricted to the vicinity of water, though adults often occur on stony rivers, brooks and upper zone of

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sea beaches, but usually found on open, moderately wet ground on peaty or gravelly ground with not too dense vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or drift. Adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Boreonebria) gyllenhali castanipes (Kirby) 1837 CHESTNUT-LEGGED GAZELLE BEETLE Helobia castanipes Kirby, 1837:20 Nebria moesta LeConte, 1850:209 Nebria elias Motschulsky, 1865:276 Nebria labradorica Casey, 1920:151 Nebria prominens Casey, 1920:151 Nebria curtulata Casey, 1924:20

Distribution: Native, New World. CANADA – AB, BC, LB, MB, NF, NT, ON, PQ, SK, YT; GREENLAND; USA – AK, ID, ME, MI, MN, MT, NH, NV, NY, OR, SD, WA, WY. Way of Life: MACROHABITAT: Lowlands to mountains, subalpine, and alpine zones, sea level – 3179 meters altitude. MICROHABITAT: On banks of cold rivers, brooks, and rivulets, as well as at the seashore. Adults occur sometimes a distance from the water in the shade of shrubs (e.g. Alnus), on rather open tundra and mountain tops on wet stony, gravelly, or silty ground mixed with sand or detritus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – November; copulating pairs in July; gravid females in late June; teneral adults in early summer and fall. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones, fallen trees and branches, and dry leaves. Adults and larvae hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Boreonebria) gyllenhali lassenensis Kavanaugh 1979 LASSEN GAZELLE BEETLE Nebria gyllenhali lassenensis Kavanaugh, 1979:96

Distribution: Native, New World. USA – CA, OR. Way of Life: M ACROHABITAT : Uplands to mountains, 2440 – 2900 meters altitude. MICROHABITAT: Hygrophilus, on lake shores and margins of brooks, on rather wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – August, and October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift or among stones. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Boreonebria) gyllenhali lindrothi Kavanaugh 1979 LINDROTH’S GAZELLE BEETLE Nebria gyllenhali lindrothi Kavanaugh, 1979:97

Distribution: Native, New World. USA – AZ, CO, NM, UT, WY. Way of Life: M ACROHABITAT : Uplands to mountains, 2130 – 3960 meters altitude. MICROHABITAT: Hygrophilus, on lake shores and margins of brooks, on rather wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September; teneral adults in July. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift or among stones. Adults hibernate. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) haida Kavanaugh 1984 HAIDA GAZELLE BEETLE Nebria haida Kavanaugh, 1984:162

Distribution: Native, New World. CANADA – BC, QC. Way of Life: MACROHABITAT: Midlands, alpine zone, 640 – 1000 meters altitude According to Kavanaugh (1992), this species is restricted to the alpine zone. MICROHABITAT: Adults and larvae are found under stones on ridges and summits above tree line, especially in areas of sparse vegetation. Unlike most alpine Nebria, members of this species avoid waterside areas and edges of snowfields and are instead most abundant in the driest sites available, moist ground with little vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Boreonebria) hudsonica LeConte 1863 HUDSONIAN BAY GAZELLE BEETLE Nebria hudsonica LeConte, 1863:3

Distribution: Native, New World. CANADA – AB, BC, MB, NT, SK, YT; USA – AK, CO, ID, MT, OR, UT, WA, WY. Way of Life: MACROHABITAT: Lowlands to mountains, subalpine, and alpine zones, 350 – 3200 meters altitude. MICROHABITAT: On barren, usually stony banks of small and large running water, not confined to really cold water. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Larval hibernation is normal. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961)

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Nebria (Catonebria) ingens ingens G. Horn 1870 CUP-FOOTED GAZELLE BEETLE Nebria ingens ingens G. Horn, 1870:98 Nebria raveni Van Dyke, 1953:102

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 3350 – 4150 meters altitude. MICROHABITAT: Granite talus slopes at the edges of glaciers, snowfields, and runoff streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Lindroth (1961) Notes: Kavanugh’s suggested name, CUP-FOOTED GAZELLE BEETLE, refers to an adaptation of the tarsomeres, highly unusal in nebrines. Nebria (Catonebria) ingens riversi Van Dyke 1925 RIVERS’ GAZELLE BEETLE Nebria ingens riversi Van Dyke, 1925:115

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 2590 – 3350 meters altitude. MICROHABITAT: Margins of glaciers, snow patches, and glacial streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) jeffreyi Kavanaugh 1984 JEFFREY’S GAZELLE BEETLE Nebria jeffreyi Kavanaugh, 1984:162

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Uplands to mountains, 2190 – 2720 meters altitude. MICROHABITAT: On banks of rivers and brooks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) kincaidi balli Kavanaugh 1979 BALL’S GAZELLE BEETLE Nebria kincaidi balli Kavanaugh, 1979:107

Distribution: Native, New World. USA – OR, WA. Way of Life: MACROHABITAT: Midlands to uplands, 980 – 1830 meters altitude. MICROHABITAT: On lake shores and near waterfalls on wet ground. DISPERSAL ABILITIES: Brachypterous, hence

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flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) kincaidi kincaidi Schwarz 1900 KINCAID’S GAZELLE BEETLE Nebria Kincaidi Kincaidi Schwarz, 1900:525 Nebria columbiana Casey, 1913:48

Distribution: Native, New World. CANADA – BC; USA – AK, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 5 – 1830 meters altitude. MICROHABITAT: On margins of forested brooks, in the shade on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. Adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) labontei Kavanaugh 1984 LABONTE’S GAZELLE BEETLE Nebria labontei Kavanaugh, 1984:163

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Mountains, 2040 – 2440 meters altitude. MICROHABITAT: Wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July and September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Boreonebria) lacustris bellorum Kavanaugh 1979 BELLS’ GAZELLE BEETLE Nebria lacustris bellorum Kavanaugh, 1979:95

Distribution: Native, New World. USA – NC, TN. Way of Life: MACROHABITAT: Midlands to uplands, 520 – 1520 meters altitude. MICROHABITAT: Moderately hygrophilus and found on banks of rivers, brooks, on shaded wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Boreonebria) lacustris lacustris Casey 1913 LACUSTRINE GAZELLE BEETLE Nebria lacustris lacustris Casey, 1913:56

Distribution: Native, New World. CANADA – MB, NB, ON, PQ; USA – CT, DC, IA, IL, IN, KY, MA, MD, ME, MI, MN, NC, NH, NJ, NY, OH, PA, TN, VA, VT, WI, WV. Way of Life: MACROHABITAT: Lowlands to uplands, 50 – 1920 meters altitude. MICROHABITAT: Near Plummers Island, Maryland, found commonly in the shade on gravel bars at the mouth of Rock Run on sand mixed with clay, under stones, at the southeast corner of Island on coarse sand mixed with silt, by splashing. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – November; teneral adults in April – May. BEHAVIOR: These beetles hide during the day and forage at night on dead and dying insects along the stream margin. Adults are nocturnal, carnivorous, taking cover in the day unders stones and wood. Hibernation takes place probably both as an adult and larva. References: Erwin (1981, 2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) lituyae Kavanaugh 1979 LITUYA’S GAZELLE BEETLE Nebria lituyae Kavanaugh, 1979:100

Distribution: Native, New World. CANADA – BC; USA – AK. Way of Life: MACROHABITAT: Midlands, alpine zone, 610 – 1100 meters altitude. MICROHABITAT: On hill summits on moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Climber on fallen trees. SEASONAL OCCURRENCE: Adults have been found in June – September; copulating pairs in August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Hibernation, at least in part, as larvae. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) louiseae Kavanaugh 1984 SOUTH MORESBY GAZELLE BEETLE Nebria louiseae Kavanaugh, 1984:162

Distribution: Native, New World. CANADA – QC. Way of Life: MACROHABITAT: Coastal lowlands, 3 – 5 meters altitude. According to Kavanaugh (1992), adults of this species are restricted to north-facing, cobble-type upper sea beaches that are at least partially shaded by the adjacent forest edge. MICROHABITAT: Stenotopic, found only where cobble is piled at least six inches deep and freshwater seeps through the sand/gravel substrate below. Some beaches on which this species occurs are strewn with driftwood. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August; copulating pairs in August. BEHAVIOR: Adults are found in great numbers climbing, running, feeding, and copulating on driftwood at night. Adults are nocturnal, carnivorous, taking cover in the day under the cobble. Hibernation, at least in part, as larvae. References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003)

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Nebria (Reductonebria) lyelli Van Dyke 1925 MOUNT LYELL GAZELLE BEETLE Nebria lyelli Van Dyke, 1925:120

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 2590 – 3510 meters altitude. MICROHABITAT: On the margins of glacial streams on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) mannerheimii Fischer von Waldheim 1828 MANNERHEIM’S GAZELLE BEETLE Nebria Mannerheimii Fischer von Waldheim, 1828:253 Nebria oregona Casey, 1913:52 Nebria corvallis Casey, 1924:20 Nebria Hippisleyi Casey, 1924:21

Distribution: Native, New World. CANADA – BC, QC; USA – AK, ID, OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1520 meters altitude. MICROHABITAT: Hygrophilus, along stony, barren margins of running streams and rivers. According to Kavanaugh (1992), on the Queen Charlotte Islands adults of this species are widespread in open areas in the lowlands, on upper sea beaches, where substrate may be sand, fine or coarse gravel, or cobble, in supra-tidal meadows, on open ground under cover (e.g. in naturally disturbed areas or synanthropic sites such as quarries), and under stones on open, exposed inorganic shores of large lowland streams. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – November; teneral adults in May – June. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Both larvae and adults hibernate. References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) meanyi giulianii Kavanaugh 1981 GIULIANI’S GAZELLE BEETLE Nebria meanyi giulianii Kavanaugh, 1981:441

Distribution: Native, New World. USA – CA, NV. Way of Life: MACROHABITAT: Mountains, 2320 – 2930 meters altitude. MICROHABITAT: On the border of brooks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Catonebria) meanyi lamarckensis Kavanaugh 1979 LAMARCK’S GAZELLE BEETLE Nebria meanyi lamarckensis Kavanaugh, 1979:109

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Uplands to mountains, 1950 – 3350 meters altitude. MICROHABITAT: On the border of brooks on wet ground.DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) meanyi meanyi Van Dyke 1925 MEANY’S GAZELLE BEETLE Nebria Meanyi Meanyi Van Dyke 1925:118

Distribution: Native, New World. CANADA – BC, YT; USA – AK, CA, OR, WA. Way of Life: MACROHABITAT: Midlands to uplands, 840 – 2190 meters altitude. MICROHABITAT: On the border of mountain streams with very cold water on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) meanyi sylvatica Kavanaugh 1979 SYLVAN GAZELLE BEETLE Nebria meanyi sylvatica Kavanaugh, 1979:109

Distribution: Native, New World. CANADA – BC; USA – WA. Way of Life: MACROHABITAT: Midlands, 670 – 1220 meters altitude. MICROHABITAT: Near springs on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) metallica Fischer von Waldheim 1820 METALLIC GAZELLE BEETLE Nebria metallica Fischer von Waldheim, 1820: pl. 6 Nebria pacifica Chaudoir, 1850:424

Distribution: Native, New World. CANADA – AB, BC; USA – AK, ID, MT, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 3 – 1830 meters altitude. MICROHABITAT: Hygrophilus, and in the southern part of its range, adults are confined to the vicinity of running water where they occur some distance from the bank under cover of shrubs and trees, sometimes on clayish ground. On the Alaskan coast and adjacent islands, adults frequent open country

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and are independent of water. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones or dead leaves. Hibernation takes place in both larval and adult forms. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) navajo Kavanaugh 1979 NAVAJO GAZELLE BEETLE Nebria navajo Kavanaugh, 1979:104

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Canyonlands of upland desert, 1940 – 2040 meters altitude. MICROHABITAT: Wet sandy ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Adults are nocturnal, carnivorous. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.) Nebria (Boreonebria) nivalis gaspesiana Kavanaugh 1979 GASPÉ GAZELLE BEETLE Nebria nivalis gaspesiana Kavanaugh, 1979:96

Distribution: Native, New World. CANADA – LB, NF, PQ; USA – ME. Way of Life: MACROHABITAT: Midlands to uplands, alpine zone, 910 – 1580 meters altitude. MICROHABITAT: On banks of cold rivers and brooks, and close to snowfields on open bare ground that is wet and stony. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – September; copulating pairs in late July; teneral adults in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Hibernation takes place probably both as an adult and larva. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Boreonebria) nivalis nivalis (Paykull) 1790 SNOW GAZELLE BEETLE Carabus nivalis Paykull, 1790:52 Nebria bifaria von Mannerheim, 1853:128 Nebria femoralis Motschulsky, 1859:541 Nebria mollis Motschulsky, 1866:274 Nebria molbis Motschulsky, 1866:274 Nebria femorata Motschulsky, 1866:275 Nebria obscuripes Poppius, 1906:19 Nebria changaica Horvatovich, 1973:251

Distribution: PALEARCTIC; CANADA – BC, NT, NU, PQ, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands to midlands, coastal tundra, and alpine zone, 25 – 1370 meters altitude. MICROHABITAT: The main habitat of this subspecies seems to be the

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margins of snowfields north of, and above tree line, but adults follow rivers and big brooks with cold water, in particular glacier outflows, deep down into the forest region. They always keep close to the water’s edge, usually on barren banks, e.g. on bare, dry sand. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Larvae, at least in part, hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) obliqua chuskae Kavanaugh 1979 CHUSKA MOUNTAINS GAZELLE BEETLE Nebria obliqua chuskae Kavanaugh, 1979:104

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands and mountainous desert, 2065 – 2320 meters altitude. MICROHABITAT: Along margins of mountain streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous. Adults take cover under stones. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.) Nebria (Reductonebria) obliqua obliqua LeConte 1866 OBLIQUE GAZELLE BEETLE Nebria obliqua LeConte, 1866:363 Nebria obtusa LeConte, 1878:478 Nebria incerta Casey, 1913:53 Nebria testaceipes Casey, 1913:54 Nebria texana Casey, 1913:54

Distribution: Native, New World. CANADA – AB, BC, NT, SK, YT; USA – AK, CA, CO, ID, MT, NE, NM, NV, OR, SD, UT, WA, WY. Way of Life: MACROHABITAT: Lowlands to mountains, 400 – 3660 meters altitude. MICROHABITAT: Markedly hygrophilus, adults occur close to the the water’s edge on the banks of larger rivers; they are not dependent on cold, clear water and are mostly found on silty places where the river runs slowly. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Larvae hibernate. Adults are nocturnal, carnivorous, taking cover in the day in cracks in the substrate. Larvae hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) ovipennis LeConte 1878 OVAL GAZELLE BEETLE Nebria ovipennis LeConte, 1878:477

Distribution: Native, New World. USA – CA, NV.

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Way of Life: MACROHABITAT: Uplands to mountains, and subalpine zone, 2040 – 3510 meters altitude. MICROHABITAT: In meadows and near snowfields on wet or moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in July – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. Hibernation takes place in both larval and adult forms. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) pallipes Say 1823 PALE-LEGGED GAZELLE BEETLE Nebria pallipes Say, 1823:78 Nebria Muehlenbergii Sturm, 1926:173

Distribution: Native, New World. CANADA – NB, NS, ON, PE, PQ; USA – CT, DC, DE, GA, IL, IN, KY, MA, MD, ME, MI, NC, NH, NJ, NY, OH, PA, RI, SC, TN, VA, VT, WI, WV. Way of Life: MACROHABITAT: Lowlands to uplands, 30 – 1520 meters altitude. MICROHABITAT: Near Plummers Island, Maryland, adults of this species are found with those of N. lacustris under stones on gravel bars at the mouth of Rock Run. Elsewhere, found on the banks of clear, swift rivers and brooks under the shade of trees and shrubs on wet, bare, stony ground. DISPERSAL ABILITIES: Wing-dimorphic, but flight of the macropterous form has not been recorded. SEASONAL OCCURRENCE: Adults have been found in March – November; copulating pairs in August; teneral adults in May – August, mostly before July. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones, wood, branches, or dead leaves. It is likely both larvae and adults hibernate. Adults are gregarious. References: Erwin (1981, 2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Boreonebria) paradisi Darlington 1931 PARADISE GAZELLE BEETLE Nebria paradisi Darlington, 1931:104 Nebria kincaidi Bänninger, 1925:264 Nebria columbiana Van Dyke, 1926:10 Nebria vandykei Darlington, 1930:24

Distribution: Native, New World. USA – OR, WA. Way of Life: MACROHABITAT: Midlands to mountains, alpine and subapline zones, 1160 – 2440 meters altitude. MICROHABITAT: Moderately hygrophilus and found on margins of streams and snowfields on rather wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Catonebria) piperi Van Dyke 1925 PIPER’S GAZELLE BEETLE Nebria piperi Van Dyke, 1925:117

Distribution: Native, New World. CANADA – BC; USA – AK, OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, and subalpine zone, 100 – 1650 meters altitude. MICROHABITAT: Under large stones near the water on barren, stony banks of rivers with cold water. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – October; tenerals in June. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Larvae hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) piute piute Erwin & Ball 1972 PIUTE GAZELLE BEETLE Nebria trifaria piute Erwin & Ball, 1972:95

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Mountains, 2510 – 3140 meters altitude. MICROHABITAT: Independent of water on loamy ground in coniferous woods near a lake. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under branches, stones, or other debris on the forest floor. References: Erwin (2001), Kavanaugh (1985, in litt.) Nebria (Catonebria) piute sevieri Kavanaugh 1984 SEVIER PLATEAU GAZELLE BEETLE Nebria piute sevieri Kavanaugh, 1984:164

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Mountains, 2440 – 3440 meters altitude. MICROHABITAT: Wet ground in forest and along edges of mountain streams under stones. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.) Nebria (Catonebria) piute utahensis Kavanaugh 1979 UTAH GAZELLE BEETLE Nebria trifaria utahensis Kavanaugh, 1979:110

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Mountains, 2290 – 2500 meters altitude. MICROHABITAT: Wet ground in forest and along edges of mountain streams under stones. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL

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OCCURRENCE:

Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.) Nebria (Catonebria) purpurata LeConte 1878 PURPLE GAZELLE BEETLE

Nebria purpurata LeConte, 1878:477 Nebria mobilis Casey, 1913:50

Distribution: Native, New World. USA – CO, NM. Way of Life: MACROHABITAT: Mountains, subalpine and alpine zones, 2350 – 3810 meters altitude. MICROHABITAT: Along cold mountain streams on wet sandy ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September; copulating pairs in June. BEHAVIOR: Adults are nocturnal, taking cover in the day under stones; they are carnivorous. They are good swimmers and often stand on rocks in the middle of the stream at night. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Reductonebria) sahlbergii modoc Kavanaugh 1979 MODOC GAZELLE BEETLE Nebria sahlbergii modoc Kavanaugh, 1979:99

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Uplands, 1740 – 1930 meters altitude. MICROHABITAT: Along brooks on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – July. BEHAVIOR : Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) sahlbergii sahlbergii Fischer von Waldheim 1828 SAHLBERG’S GAZELLE BEETLE Nebria Sahlbergii Fischer von Waldheim, 1828:254 Nebria violacea Motschulsky, 1850:73 Nebria aleuta Van Dyke, 1924:5

Distribution: Native, New World. CANADA – AB, BC, NT, QC, YT; USA – AK, MT, OR, WA. Way of Life: MACROHABITAT: Lowlands to mountains, and alpine zone, 3 – 2530 meters altitude, MICROHABITAT: Along streams, sometime very small ones, lake shores at high altitudes on shady wet gravelly-stony ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – August; teneral adults in March – November. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Likely both adults and larvae hibernate. References: Erwin (2001), Kavanaugh (1985, 1992, in litt.), Larochelle & Larivière (2003)

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Nebria (Reductonebria) sahlbergii triad Kavanaugh 1979 TRINITY GAZELLE BEETLE Nebria sahlbergii triad Kavanaugh, 1979:99

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Midlands to uplands, 910 – 1740 meters altitude. MICROHABITAT: Along stream margins on wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Found Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) schwarzi beverlianna Kavanaugh 1979 BEVERLY ANNE’S GAZELLE BEETLE Nebria schwarzi beverlianna Kavanaugh, 1979:110

Distribution: Native, New World. USA – WY. Way of Life: MACROHABITAT: Uplands, 2100 meters altitude. MICROHABITAT: Along river banks on gravelly ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) schwarzi schwarzi Van Dyke 1925 SCHWARZ’S GAZELLE BEETLE Nebria Schwarzi Van Dyke, 1925:116

Distribution: Native, New World. CANADA – AB, BC. Way of Life: MACROHABITAT: Midlands, 1100 – 1430 meters altitude. MICROHABITAT: Along stream margins on wet bare coarse sand. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. Adults hibernate. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) sierrablancae Kavanaugh 1984 SIERRA BLANCA GAZELLE BEETLE Nebria sierrablancae Kavanaugh, 1984:164

Distribution: Native, New World. USA – NM. Way of Life: MACROHABITAT: Mountains, 2740 – 3510 meters altitude. MICROHABITAT: Shaded wet ground in forests and under stones along edges of small shaded streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.)

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Nebria (Catonebria) spatulata sierrae Kavanaugh 1979 SIERRAN GAZELLE BEETLE Nebria spatulata sierrae Kavanaugh, 1979:108

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, alpine zone, 2740 – 3480 meters altitude. MICROHABITAT: On brook margins and lake shores on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift or stones. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) spatulata spatulata Van Dyke 1925 SPATULATE GAZELLE BEETLE Nebria spatulata Van Dyke, 1925:119 Nebria fusiformis Van Dyke, 1926:11

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, alpine zone, 3050 – 3140 meters altitude MICROHABITAT: On brook margins and lake shores on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift or stones. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) steensensis Kavanaugh 1984 STEENS MOUNTAINS GAZELLE BEETLE Nebria steensensis Kavanaugh, 1984:165

Distribution: Native, New World. USA – OR. Way of Life: M ACROHABITAT : Uplands to mountains, 2190 – 2725 meters altitude. MICROHABITAT: Margins of small streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – September. BEHAVIOR: Adults are nocturnal, taking cover in the day; they are carnivorous. References: Erwin (2001), Kavanaugh (1985, in litt.) Nebria (Reductonebria) suturalis LeConte 1850 SEAMED GAZELLE BEETLE Nebria suturalis LeConte, 1850:209 Nebria longula LeConte, 1878:478 Nebria nimbosa Casey, 1920:150

Distribution: Native, New World. CANADA – AB, LB, ON, PQ; USA – CO, NH, NY, VT, WY.

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Way of Life: MACROHABITAT: Midlands to mountains, subalpine, and alpine zones, 1190 – 4350 meters altitude. MICROHABITAT: Confined to mountain tops; adults are found under stones where water is seeping or trickling down the mountain side in open places above tree line. Some individuals may occur a little below tree line, along brooks. DISPERSAL ABILITIES: Wing-dimorphic, neither form capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in late July – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones, larvae in crevices. Adults hibernate. References: Erwin (2001), Bell (1978), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Catonebria) trifaria pasquineli Kavanaugh 1984 PASQUINEL’S GAZELLE BEETLE Nebria trifaria pasquineli Kavanaugh, 1984:165

Distribution: Native, New World. USA – CO, WY. Way of Life: MACROHABITAT: Mountains, 2290 – 3960 meters altitude. MICROHABITAT: Near streams and lakes. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) trifaria trifaria LeConte 1878 NORTHERN CHAIN-BACKED GAZELLE BEETLE Nebria trifaria LeConte, 1878:478 Nebria tetonensis Erwin & Ball, 1972:95

Distribution: Native, New World. USA – ID, MT, NV, UT, WY. Way of Life: M ACROHABITAT : Uplands to mountains, 1680 – 3470 meters altitude. MICROHABITAT: Near snowfields on wet ground and along margins of mountain streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003), Lindroth (1961) Nebria (Boreonebria) turmaduodecima Kavanaugh 1981 TROUP TWELVE’S GAZELLE BEETLE Nebria turmaduodecima Kavanaugh, 1981:436

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Uplands to mountains, alpine zone, 2100 – 2290 meters altitude. MICROHABITAT: Margins of snowfields. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in day under stones. References: Erwin (2001), Kavanaugh (1981, 1985, in litt.), Larochelle & Larivière (2003)

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Nebria (Catonebria) vandykei vandykei Bänninger 1928 VAN DYKE’S GAZELLE BEETLE Nebria Vandykei Vandykei Bänninger, 1928:5

Distribution: Native, New World. USA – WA. Way of Life: MACROHABITAT: Midlands to uplands, subalpine and alpine zones, 1100 – 2040 meters altitude. MICROHABITAT: On stream margins on wet ground and at edges of snowfields. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August, and October. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Catonebria) vandykei wyeast Kavanaugh 1979 WYEAST’S GAZELLE BEETLE Nebria vandykei wyeast Kavanaugh, 1979:109

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Uplands, subalpine and alpine zones, 1680 – 2190 meters altitude. MICROHABITAT: On stream margins on wet ground and at edges of snowfields. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1979, 1985, in litt.) Nebria (Reductonebria) wallowae Kavanaugh 1984 WALLOWA MOUNTAINS GAZELLE BEETLE Nebria wallowae Kavanaugh, 1984:161

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Uplands to mountains, 1980 – 2440 meters altitude. MICROHABITAT: On lake shores and stream margins, and in meadows. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July and September. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under drift and stones. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) zioni oasis Kavanaugh 1979 OASIS GAZELLE BEETLE Nebria zioni oasis Kavanaugh, 1979:103

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Uplands, 1920 – 1980 meters altitude. MICROHABITAT: On brook margins. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day.

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References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nebria (Reductonebria) zioni zioni Van Dyke 1943 ZION GAZELLE BEETLE Nebria zioni Van Dyke, 1943:20

Distribution: Native, New World. USA – UT. Way of Life: MACROHABITAT: Midlands to mountains, often in canyons, 1360 – 3120 meters altitude. MICROHABITAT: On brook margins. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1985, in litt.), Larochelle & Larivière (2003) Nippononebria Uéno 1955 FALSE GAZELLE BEETLES (Plate 6) Nippononebria Uéno, 1955:49 Vancouveria Kavanaugh, 1995:156 [valid subgenus for native Nearctic species]

Number of worldwide species: Seven. Number of Western Hemisphere species: Three. Taxonomy: Stable. Adelphotaxon: Leistus Frölich 1799 Type species: Nebria pusilla Ueno, 1955. Distribution: Native, New World (three species), Palearctic (four species). Habitat: Wet meadows, at low to high altitudes. References: Erwin (2001), Kavanaugh (1995) Nippononebria (Vancouveria) altisierrae (Kavanaugh) 1984 HIGH SIERRA FALSE GAZELLE BEETLE Nebria altisierrae Kavanaugh, 1984:160

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Mountains, 2560 – 3110 meters altitude, in meadows. MICROHABITAT: Wet ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – August, and November. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. References: Erwin (2001), Kavanaugh (1984, in litt.), Larochelle & Larivière (2003)

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Nippononebria (Vancouveria) campbelli (Kavanaugh) 1984 CAMPBELL’S FALSE GAZELLE BEETLE Nebria campbelli Kavanaugh, 1984:161

Distribution: Native, New World. CANADA – BC; USA – WA. Way of Life: MACROHABITAT: Uplands, 1520 – 2130 meters altitude. MICROHABITAT: According to Kavanugh (in litt.), well above treeline at Mount Baker, under a stone on a rocky outcrop surrounded by huge snowfields on one of the mountain’s main ridges. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day under stones. References: Erwin (2001), Kavanaugh (1984, in litt.), Larochelle & Larivière (2003) Nippononebria (Vancouveria) virescens (G. Horn) 1870 GREENISH FALSE GAZELLE BEETLE Nebria virescens G. Horn, 1870:10 Nebria brevis Casey, 1913:55

Distribution: Native, New World. CANADA – BC; USA – CA, ID, OR, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 230 – 1680 meters altitude, meadows. MICROHABITAT: Independent of water, on rather dry, slightly shaded meadow substrate. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – December. BEHAVIOR: Adults are nocturnal, carnivorous, taking cover in the day. Adults, at least in part, hibernate. References: Erwin (2001), Kavanaugh (in litt.), Larochelle & Larivière (2003), Lindroth (1961) Notiokasiini Kavanaugh & Nègre 1983 SOUTHERN SISTER BEETLES Notiokasiini Kavanaugh & Nègre, 1983:551

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Notiophilus Dumeril 1806 + Opisthini Geographic status: Neotropical Distribution: Native, New World. Brazil; Uruguay. References: Erwin (2001), Kavanaugh & Nègre (1983) Notiokasis Kavanaugh & Nègre 1983 SOUTHERN SISTER BEETLES (Plate 7) Notiokasius, Erwin, 1979:591 [nomen nudum] Notiokasis Kavanaugh & Nègre, 1983:551

Number of worldwide species: One. Number of Western Hemisphere species: One.

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Taxonomy: Stable. Adelphotaxon: See Notiokasiini, above. Type species: Notiokasis chaudoiri, Kavanaugh & Nègre, 1983 Distribution: Native, New World. Brazil; Uruguay. (One species). Habitat: Lowlands. Temperate maritime climate with warm summers, cool winters and 100 – 150 mm annual rainfall. Microhabitat unknown. References: Erwin (2001), Kavanaugh & Nègre (1983) Notiokasis chaudoiri Kavanaugh & Nègre 1983 CHAUDOIR’S SOUTHERN SISTER BEETLE Notiokasis chaudoiri Kavanaugh & Nègre, 1983:558

Distribution: Native, New World. Brazil; Uruguay. Way of Life: MACROHABITAT: Humid subtropical lowlands. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Unknown. References: Erwin (2001), Kavanaugh & Nègre (1983) Opisthiini Dupuis 1912 FALSE MARSH AND BOG BEETLES Opisthiini Dupuis 1912:1

Number of worldwide genera: Two. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Notiokasini + Notiophilini Geographic status: Holarctic. References: Erwin (2001), Lindroth (1961), Kavanaugh (1996, 1999) Opisthius Kirby 1837 FALSE MARSH AND BOG BEETLES (Plate 8) Opisthius Kirby, 1837:59

Number of worldwide species: One. Number of Western Hemisphere species: One. Taxonomy: Stable. Adelphotaxon: Paropisthius Casey 1920 Type species: Opisthius richardsoni Kirby, 1837 Distribution: Native, New World (one species). Habitat: Banks of rivers and big brooks. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961)

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Opisthius richardsoni Kirby 1837 RICHARDSON’S FALSE MARSH AND BOG BEETLE Opisthius Richardsoni Kirby, 1837:61

Distribution: Native, New World. CANADA – AB, BC, NT, SK, YT; USA – AK, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY. Way of Life: MACROHABITAT: Lowlands to mountains, 70 – 2652 meters altitude. MICROHABITAT: Confined to the banks of rivers and big brooks, on places where the substrate is soft, consisting of clay, often mixed with sand and gravel. DISPERSAL ABILITIES: Macropterous, capable of flight, frequent flyer. SEASONAL OCCURRENCE: Adults have been found in February – October; teneral adults in July – August. BEHAVIOR: Larvae dig burrows, as a rule in the company of the Staphylinds of the genus Bledius. Adults keep close to the water where the ground is free of vegetation. They have been observed flying in the sunshine. Adults hibernate. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Notiophilini Mostschulsky 1850 BIG-EYED BEETLES Notiophilini Mostschulsky, 1850:16

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Opisthini Dupuis 1912 Geographic status: Holarctic, Neotropical. References: Erwin (2001), Barsevskis (1994), Lindroth (1961) Notiophilus Dumeril 1806 BIG-EYED BEETLES (Plate 9) Notiophilus Dumeril, 1806:193 Nothiophilus Dumeril, 1806:194 Nothiophilus Say, 1823:81 Latviaphilus Barsevskis, 1994:1 Makarovius Barsevskis, 1994:1

Number of worldwide species: 54. Number of Western Hemisphere species: 18. Taxonomy: Stable. Adelphotaxon: See Notiophilini, above. Type species: Cicindela aquatica Linné, 1758. Distribution: Native, New World (14 species), Introduced (two species), Holarctic (two species), Palearctic (28 species). Habitat: According to Lindroth (1961), members of this genus are heliophorus and rather xerophilus. Most species occur in open country where the vegetation is sparse and the ground consists of gravel, others belong to open forested areas. Adults are pronouncedly carnivorous,

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seen feeding on gants and spiders. Wing dimorphism occurs more regularly in this genus than any other large carabid genus. References: Erwin (2001), Barsevskis (1994), Lindroth (1961) Notiophilus aeneus (Herbst) 1806 BRASSY BIG-EYED BEETLE Elaphrus aeneus Herbst, 1806:235 Notiophilus porrectus Say, 1830: (3) [3]

Distribution: Native, New World. CANADA – NB, NS, ON, PE, PQ; USA – CT, DC, DE, GA, IA, IL, IN, MA, MD, ME, MI, MN, MO, NC, NH, NJ, NY, OH, PA, RI, SC, TN, VA, VT, WI, WV. Way of Life: MACROHABITAT: Lowlands, 3 – 335 meters altitude, in deciduous forests (e.g. Acer, Quercus, Ostrya, mixed forests, and coniferous forests (e.g. Pinus); forest clearings and roadsides. MICROHABITAT: At Plummers Island, Maryland, adults were found in moss and deep leaf litter near damp places in deciduous woods, especially near the Trail Pond. DISPERSAL ABILITIES: Macropterous, adults occasionally found at lights at night. Climbers on the bases of trees. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Adults are mostly diurnal and active in the sunshine; they are carnivorous, probably predaceous on small arthropods and were recorded feeding on lepidopterous larvae. Adults hibernate; found over-wintering in woods and at the edge of woods. References: Erwin (1981, 2001), Larochelle (1975), Larochelle & Larivière (2003), Lindroth (1961), (Reeves, et al. 1983), Data from CAS, NMNH, UASM collections Notiophilus aquaticus (Linné) 1758 LINNÉ’S BIG-EYED BEETLE Cicindela aquaticus Linné, 1758:408 Cicindela pusillus Schreber, 1759:10 Elaphrus semipunctatus Fabricius, 1775:227 Notiophilus metallicus Waterhouse, 1833:203 Notiophilus parvulus Waterhouse, 1833:208 Notiophilus dauricus Motschulsky, 1860:539 Notiophilus strigifrons Baudi di Selve, 1864:146 Notiophilus Hardyi Putzeys, 1866:165 Notiophilus apicalis Dalla Torre, 1877:22 Notiophilus obscurus Dalla Torre, 1877:22 Notiophilus obsoletus Dalla Torre, 1877:22 Notiophilus Dybowskii Lomnicki, 1903:113 Notiophilus Kroli Lomnicki, 1903:113 Notiophilus pristinus Friederichs, 1903:88 Notiophilus Semenovi Tschitschérine, 1903:116 Notiophilus Blacki Edwards, 1913:70 Notiophilus Spaethi Munster, 1922:140 Notiophilus aquaticus bimaculatus Barševskis, 1993:51 Notiophilus aquaticus pseudoaesthuans Barševskis, 1994:7

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Notiophilus aquaticus stoschkae Barševskis, 1994:7 Notiophilus fuscipes Barševskis, 1999:79

Distribution: Holarctic. PALEARCTIC; CANADA – AB, BC, LB, MB, NB, NF, NS, ON, PM, PQ, SK, YT; USA – AK, AZ, CO, IL, MA, ME, MI, MN, MT, NH, NM, NY, PA, VT, WI. Way of Life: MACROHABITAT: Lowlands to mountains, subalpine, alpine zones, and coastal tundra, sea level – 3048 meters altitude. MICROHABITAT: On moraines, fields, meadows, vacant lots, cultivated fields (e.g. corn, barley), roadsides, sand pits, gravel pits, forest edges, and in mixed deciduous forests. Adults occur on open ground that is well-drained, and consisting of moderately dry gravelly, or sandy ground covered with sparse or moderate vegetation. Gandhi, et al. (2005) reported them also from Jack Pine (Pinus banksiana Lamb.) wind-disturbed prescribedburned forest. DISPERSAL ABILITIES: Wing-dimorphic. European records indicate macropterous adults fly. SEASONAL OCCURRENCE: Adults have been found in January – December; teneral adults in June – August. BEHAVIOR: In Europe, adults have been observed eating Collembola. Adults are mostly diurnal, carnivorous. Adults found over-wintering on higher and drier ground, in fields and at the edge of woods under stones and in leaf litter. Larvae and adults hibernate. References: Erwin (2001), Gandhi et al. (2005), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, UASM collections Notiophilus biguttatus (Fabricius) 1779 SPOTTED BIG-EYED BEETLE Elaphrus biguttatus Fabricius, 1779:232 Dermestes aquaticus Geoffroy, 1785:21 Elaphrus semipunctatus Duftschmid, 1812:90 Notiophilus latus Waterhouse, 1833:209 Notiophilus nitidus Waterhouse, 1833:210 Notiophilus striatus Waterhouse, 1833:209 Notiophilus lateralis Motschulsky, 1864:192 Notiophilus melanophthalmus Slosser-Klekovski, 1877:12 Notiophilus pseudoquadripunctatus Everts, 1918:43 Notiophilus pseudolateralis Louvet, 1925:6 Notiophilus coerulescens Depoli, 1929:31

Distribution: Introduced. PALEARCTIC; CANADA – BC, NB, NF, NS, PM; USA – ME, NH. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 900 meters altitude. In Europe, this species is a pronounced forest beetle except in the humid climate of the west, where it occurs in open country. MICROHABITAT: Clearings, roadsides, vacant lots, open grassy cultivated fields, and forest edges. Adults occur on shaded or open ground that is consisting of moderately dry gravelly, sandy, or loamy ground covered with sparse vegetation, or leaf litter. DISPERSAL ABILITIES: Wing-polymorphic. European records indicate macropterous adults fly. SEASONAL OCCURRENCE: Adults have been found in May – November; teneral adults in June – September. BEHAVIOR: Adults are mostly diurnal, taking cover on cloudy days under dead leaves or stones, among conifer needles or pieces of wood; they are carnivorous and in Europe, adults have been observed eating Collembola, mites, flies, and other arthropods. Larvae are known to climb on tree trunks. This species is favored by human activities and is an effective colonist. Adults hibernate. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, UASM collections

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Notiophilus borealis Harris 1869 NORTHERN BIG-EYED BEETLE Notiophilus borealis Harris, 1869:213

Distribution: Holarctic. PALEARCTIC; CANADA – AB, BC, LB, MB, NF, NT, ON, PQ, SK, YT; USA – AK, ME, NH, NY, VT. Way of Life: MACROHABITAT: Lowlands to mountains, subalpine and alpine zones, 11 – 2332 meters altitude. MICROHABITAT: On open or slightly shaded rather dry, gravelly or sandy ground covered with sparse or moderately short vegetation (e.g., mosses) on moraines, or in meadows, vacant lots, along roadsides, in forest clearings, or open forests and above timber line on Mt. Washington in NH. DISPERSAL ABILITIES: Wing-dimorphic: brachypterous or macropterous. Macropterous individuals probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – October; tenerals in August – September. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days under stones and in moss. Adults are solitary. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, UASM collections Notiophilus chihuahuae Casey 1913 CHIHUAHUA BIG-EYED BEETLE Notiophilus chihuahuae Casey, 1913:47

Distribution: Native, New World. MÉXICO – CH, DU, SI, SO. Way of Life: MACROHABITAT: Uplands to mountains, 2100–2820 meters altitude, in oak/pine forests and at higher altitudes in oak/pine/fir forests. MICROHABITAT: In leaf litter, particularly conifer litter. DISPERSAL ABILITIES: Brachypterous (at least in part), hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – September. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days in litter. References: Erwin (2001), Data from NMNH, UASM collections Notiophilus directus Casey 1920 STRAIGHT BIG-EYED BEETLE Notiophilus directus Casey, 1920:142 Notiophilus lanei Hatch, 1949:114

Distribution: Native, New World. CANADA – AB, BC; USA – AZ, CA, CO, ID, MT, NM, OR, UT, WA, WY. Way of Life: MACROHABITAT: Midlands to mountains, 1500 – 3200 meters altitude. MICROHABITAT: On open bare dry, gravelly (sometimes sand-mixed) ground, or that shaded by shrubs or trees, or, covered with tiny mosses on moraines; or in the vicinity of brooks and lakes. DISPERSAL ABILITIES: Wing-dimorphic: macropterous or brachypterous. Macropterous individuals probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – November; teneral adults in July – August. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days among gravel or under pieces of wood. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH, UASM collections

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Notiophilus intermedius Lindroth 1954 INTERMEDIATE BIG-EYED BEETLE Notiophilus intermedius Lindroth, 1954:157

Distribution: Native, New World. CANADA – BC, LB, MB, NF, PQ; USA – AK. Way of Life: MACROHABITAT: Lowlands, sea level – 230 meters altitude. MICROHABITAT: Xerophilus, on open well-drained, usually dry, sandy ground covered with sparse vegetation on river banks. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – July, and September. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days under mosses, lichens, or low plants, e.g. Empetrum (Ericaceae). References: Erwin (2001), Larochelle & Larivière (2003) Notiophilus nemoralis Fall 1906 WOODLAND BIG-EYED BEETLE Notiophilus nemoralis Fall, 1906:88

Distribution: Native, New World. CANADA – [PQ]; USA – MA, ME, NH, NY, VT. Way of Life: MACROHABITAT: Midlands, 900 – 1100 meters altitude, in open coniferous forests (Abies, Picea). MICROHABITAT: On open dry ground with bare spots surrounded by vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adult in August. BEHAVIOR: Mostly diurnal; active in the sunshine among conifer needles; adults take cover on cloudy or cool days in needle litter, among dead leaves, and in moss. Adults are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Notiophilus nitens LeConte 1857 BRIGHT BIG-EYED BEETLE Notiophilus nitens LeConte, 1857:31

Distribution: Native, New World. CANADA – BC; USA – ID, MT, OR, WA. Way of Life: MACROHABITAT: Lowlands, 41 – 292 meters altitude. MICROHABITAT: On open dry, sandy, gravelly or rocky ground covered with sparse vegetation in grasslands, pastures, cultivated fields, and hills. DISPERSAL ABILITIES: Wing-dimorphic: brachypterous or macropterous. SEASONAL OCCURRENCE: Adults have been found in March – July, October; teneral adults in May – June. BEHAVIOR: Mostly diurnal, xerophilus, active in the sunshine; adults take cover on cloudy or cool days under dead fallen plants or dry cattle dung. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections Notiophilus novemstriatus LeConte 1848 NINE-LINED BIG-EYED BEETLE Notiophilus novemstriatus LeConte, 1848:350 Notiophilus cribrilaterus Motschulsky, 1864:193

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Notiophilus quadrifoveolatus T.W. Harris, 1869:212 Notiophilus parvus Casey, 1920:142

Distribution: Native, New World. CANADA – NS; USA – AL, AR, AZ, CT, DC, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, MS, MT, NC, NE, NH, NJ, NM, NY, OH, OK, PA, SC, SD, TN, TX, VA, VT. Way of Life: MACROHABITAT: Lowlands, 5 – 50 meters altitude. MICROHABITAT: Open dry ground, especially with green moss in sparse or open forests (deciduous, mixed, coniferous) near streams and ponds. Also along forest edges and in thickets. DISPERSAL ABILITIES: Wingdimorphic: brachypterous or macropterous. According to Lindroth (1961) macropterous individuals are rare. Macropterous adults occasionally found at lights at night, thus they do fly. Occasional climber on trees. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Diurnal, active in the sunshine on moss, around stones and other debris; adults take cover on cloudy or cool days under mosses and dead leaves, sticks, or bunches of grass. Adults are gregarious. Adults found overwintering among moss, dead leaves and sticks. Adult are predaceous on dipterans (e.g. gnats). References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections Notiophilus palustris (Duftschmid) 1812 MARSH BIG-EYED BEETLE Elaphrus palustris Duftschmid, 1812:192 Notiophilus nitidulus Waterhouse, 1833:203 Notiophilus parallelus Waterhouse, 1833:204 Notiophilus newmanni Waterhouse, 1833:205 Notiophilus Davisi Waterhouse, 1833:205 Notiophilus brevis Waterhouse, 1833:206 Notiophilus tibialis Waterhouse, 1833:206 Notiophilus latior Waterhouse, 1833:207 Notiophilus melancholicus Dalla Torre, 1877:23 Notiophilus hirticollis Chaudoir, 1882: LXXX Notiophilus atavus Friedrichs, 1903:88 Notiophilus decemstriatus Roubal, 1948:45 Notiophilus impunctatus Pulpan, 1948:49 Notiophilus mlynari Pulpan, 1948:49

Distribution: Holarctic. PALEARCTIC; CANADA – NS, PE. Way of Life: MACROHABITAT: Midlands to uplands, 549 – 1981 meters altitude. MICROHABITAT: On open moderately moist sandy ground covered with some vegetation. At edges of cultivated fields and in Europe in drier zones of swamps (Alnus) covered with dense tall vegetation. DISPERSAL ABILITIES: Macropterous, probably capable of flight, e.g. frequent in seashore drift material. SEASONAL OCCURRENCE: Adults have been found in Canada in June – July; teneral adults in July. In Europe, May – July, and October. BEHAVIOR: Diurnal, active in the sunshine. Adult are predaceous on springtails and mites in Europe. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections

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Notiophilus rufipes Curtis 1829 RUFUS-LEGGED BIG-EYED BEETLE Notiophilus rufipes Curtis, 1829:254 Notiophilus fulvipes Motschulsky, 1845:12 Notiophilus rufipes Chaudoir, 1844:439 Notiophilus femoralis Lomnicki, 1903:107

Distribution: Introduced. PALEARCTIC; USA – GA. Way of Life: MACROHABITAT: Lowlands to uplands, 44 – 2000 meters altitude, in mixed forests, at the forest edges, in vine and fruit tree orchards. MICROHABITAT: Usually found on the ground near the base of separately growing trees. DISPERSAL ABILITIES: Wing-dimorphic, mostly macropterous, capable of flight. SEASONAL OCCURRENCE: In Europe, March, July – August, October. BEHAVIOR: Adults have been observed hibernating in Europe. Adults move fast by running and flying, hence most are macropterus. References: Erwin (2001), Barševskis (2004), Larochelle & Larivière (2003), Data from NMNH collection Notes: The single know specimen from Georgia was collected in August of 1975 at 44 meters altitude in the Okefenokee Swamp. Whether the species has become established is unknown. Notiophilus semiopacus Eschscholtz 1833 SEMIOPAQUE BIG-EYED BEETLE Notiophilus semiopacus Eschscholtz, 1833:25

Distribution: Native, New World. MÉXICO – CH, SO; USA – AZ, CA. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 2090 meters altitude, in pine/ oak forests. MICROHABITAT: On open ground on ridges and in the vicinity of snowfields and seeps, and in foothill canyons along streams, in parks, and on the sea beach in southern CA; in México in pine litter and meadows of pine/oak forests. DISPERSAL ABILITIES: Wing-dimorphic: brachypterous or macropterous. Macropterous individuals capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days at the base of stones and in ant nests. Macropterous adults occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Notiophilus semistriatus Say 1823 SEMI-STRIATE BIG-EYED BEETLE Notiophilus semistriatus Say, 1823:81 Notiophilus confusus LeConte, 1848:349 Notiophilus punctatus LeConte, 1850:210 Notiophilus americanus T.W. Harris, 1869:213 Notiophilus coloradensis Casey, 1920:141

Distribution: Native, New World. CANADA – AB, BC, LB, MB, NS, ON, PQ, YT; USA – AK, AR, CO, CT, DC, DE, GA, IA, IL, IN, KS, KY, MA, MD, MI, MN, MO, NC, NH, NJ, NM, NY, OH, OK, PA, SC, SD, VA, WV, WY.

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Way of Life: MACROHABITAT: Lowlands to mountains, subalpine and alpine zones, 200 – 3200 meters latitude. MICROHABITAT: On open or half-shaded rather dry, gravelly, sandy, rocky or peaty ground; covered with sparse low thin vegetation or patches of mosses and lichens on moraines, in abandoned fields, or fence rows, roadsides, orchards; forest edges, and clearings. DISPERSAL ABILITIES: Wing-dimorphic: macropterous or brachypterous. Occasional climber on trees. SEASONAL OCCURRENCE: Adults have been found in January – December; teneral adults in May – June. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days under reindeer moss (Cladonia), stones, dead leaves, and in clumps of broom sedges. Adults are gregarious. Adults found over wintering under mosses and dead leaves. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Notiophilus sierranus Casey 1920 MOUNTAIN BIG-EYED BEETLE Notiophilus sierranus Casey, 1920:140 Notiophilus obscurus Fall, 1901:207 Notiophilus obscuratus Fall, 1926:125

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Midlands, 550 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Notiophilus simulator Fall 1906 FALL’S BIG-EYED BEETLE Notiophilus simulator Fall, 1906:86 Notiophilus evanescens Casey, 1913:47

Distribution: Native, New World. CANADA – AB, BC, SK, NT, YT; USA – AK, CO, ID, MT, OR, SD, TX, UT, WA, WI, WY. Way of Life: M ACROHABITAT : Uplands to mountains, 1520 – 3300 meters altitude. MICROHABITAT: On open or slightly shaded sandy bare ground that is nearly bare except for patches of the tiny moss, Ceratodon purpureus (Hedw.) Brid., and under shrubby trees (Alnus). DISPERSAL ABILITIES: Wing-dimorphic: brachypterous or macropterous. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Notiophilus specularis Bates 1881 MIRROR-WINGED BIG-EYED BEETLE Notiophilus specularis Bates, 1881:19

Distribution: Native, New World. EL SALVADOR; GUATEMALA; MÉXICO – CH, CL, DF, GO, HD, MX, MH, OA, PU, QT, SL, TL, VC.

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Way of Life: MACROHABITAT: Uplands to mountains, 1540 – 3100 meters altitude, in mesophylic and cloud forests, oak, pine/oak forests, pine/alder/manzanita forest edge, and in Avocado orchard. MICROHABITAT: On the ground at the base of trees and in leaf litter. DISPERSAL ABILITIES: Macropterous and brachypterous, macropterous likely capable of flight. SEASONAL OCCURRENCE: Adults have been found in January, April – November; teneral adults in April in CH. BEHAVIOR: Running on leaf litter at night in HD. Adults take cover under stones and in coniferous litter. References: Erwin (1991, 2001), Data from NMNH, UASM collections Notiophilus sylvaticus Eschscholtz 1833 FOREST BIG-EYED BEETLE Notiophilus sylvaticus Eschscholtz 1833:24

Distribution: Native, New World. CANADA – BC, QC; USA – AK, CA, ID, NM, OR, WA Way of Life: MACROHABITAT: Lowlands to uplands, subalpine and alpine zones, sea level – 1829 meters altitude. MICROHABITAT: On open slightly shaded gravelly ground in open forests (e.g. Pinus) and on meadows. According to Lindroth (1961), on the mainland they occur on slightly shaded, gravelly ground, but not in dense forests. Also, occasionally found in Greenhouses. According to Kavanaugh (1992), this species is widespread on the Queen Charlotte Islands in forested areas at all elevations, from near sea level to above tree line on the mountain peaks. Adults are found under the loose bark of fallen trees or in leaf litter, under stones or fallen trees on the ground, or in the deep moss layer that covers much of the forest floor throughout the archipelago. Above tree line, beetles occur in the driest sites with very low, scattered vegetation, and they are especially active in sunlight. DISPERSAL ABILITIES: Wing-dimorphic: brachypterous or macropterous, both forms occurring in the same population. Macropterous individuals probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February, and April – October; teneral adults in June and August. BEHAVIOR: Mostly diurnal, active in the sunshine; adults take cover on cloudy or cool days under the loose bark of fallen trees, under fallen trees and stones, in moss, and among conifer needles. Adults are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections CICINDITAE Cicindini Cicindini Csiki 1927 Cicindini Csiki, 1927:425

Number of worldwide genera: Two. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon unknown, but possibly Omophron Latreille, 1802. Geographic status: Palearctic, Neaustral. References: Erwin (2001), Kavanaugh & Erwin (1991), Kavanaugh (1999)

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Cicindis Bruch 1908 FAIRY SHRIMP HUNTING BEETLES (Plate 10) Cicindis Bruch, 1908:496

Number of species: One. Taxonomy: Stable. Adelphotaxon: Archaeocindis Kavanaugh & Erwin, 1991. Type species: Cicindis horni Bruch, 1908. Distribution: Native, New World. ARGENTINA – Catamarca, Córdoba Provinces. (one species). Habitat: The single known species is restricted to vernal pools on inland salt flats. References: Erwin (2001), Kavanaugh & Erwin (1991), Erwin & Aschero (2004) Cicindis horni Bruch 1908 HORN’S FAIRY SHRIMP HUNTING BEETLE Cicindis Horni Bruch, 1908:499

Distribution: Native, New World. ARGENTINA – Catamarca, Córdoba. Way of Life: MACROHABITAT: Lowlands, 150 – 300 meters altitude, margins of large salt flats near vernal pools. MICROHABITAT: Adults occur on wet to inundated clay mixed with salt where polygons form when the clay dries out and with no vegetation cover. DISPERSAL ABILITIES: Adults are fully winged and fly readily at night; adults also are swimmers and divers in vernal pools with 17% salinity. SEASONAL OCCURRENCE: Adults are found in December – March; copulating pairs observed in March; teneral adults in February. BEHAVIOR: This heretofore very rare species was recently found to be markedly unusual among carabid beetles because adults surface-swim and dive in search of their prey, an anostracan fairy shrimp, possibly a species of Thamnocephalus Cohen. Adult beetles do not run along the shore at the margin of the ponds that are formed in the great salt flats like most carabid in this type of habitat elsewhere. However, they do rest in small burrows during the day away from the water’s edge, and emerge before nightfall (crepuscular behavior). They then spend the night in search of food or mates swimming on the water surface of markedly alkaline water bodies (17% salinity). Tibial natatorial setae, tarsal setal fringes, ventral vestiture, and genal/pronotal planing surfaces plus decumbent and very sharply pointed mandibles are specific adaptations that permit this species to behave as a swimming superspecialist predator. Subsequent studies have shown that adults also run on very wet and sticky mud at the edge of vernal pools in search of mates. Mating can take place on the water surface, or on adjacent mud. Adults are best collected by net at night over the pool, by hand when they are running on the mud adjacent to the pool, or at UV and Mercury Vapor lights; and they sometimes can be obtained in pitfall traps placed very near the water’s edge. Immature stages and hibernation site unknown. References: Erwin (2001), Kavanaugh & Erwin (1991), Erwin & Aschero (2004), Data from NMNH collection

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OMOPHRONITAE Omophronini Bonelli 1810 ROUND SAND BEETLES Omophronini Bonelli, 1810: tab syn

Number of worldwide genera: Two. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Unknown, but possibly Cicindini Csiki 1927. Geographic status: Holarctic, Oriental, Afrotropical. References: Erwin (2001), Jeannel (1946) Omophron Latreille 1802 ROUND SAND BEETLES (Plate 11) Omophron Latreille, 1802:89 Scolytus Fabricius, 1790:221 Epactius Schneider, 1791:23 Lithophilus Schneider, 1791:23 Homophron Fischer von Walheim, 1828:256 Epacter Semenov, 1922:38 Andromophron Semenov, 1922:40 Oligomophron Semenov, 1922:40 Paromophron Semenov, 1922:40 Homophron Semenov, 1922:41 Stenomophron Semenov, 1922:42 Istor Semenov 1922:43 Prosecon Semenov, 1922:44 Phromoon Lutshnik, 1933:132

Number of worldwide species: 65. Number of Western Hemisphere species: 17. Taxonomy: Stable. Adelphotaxon: Phrator Semenov 1922. Type species: Carabus limbatus Fabricius, 1776. Distribution: Native, New World (17 species), Afrotropical (12 species), Nearctic (12 species), Neotropical (six species), Oriental (18 species), Palearctic (one species), two species are found in both Nearctic and Neotropical Realms. Habitat: According to Lindroth (1961), all species of this genus are markedly hygrophilous and restricted to the immediate vicinity of water where the ground is sand or clay and is bare, or with quite sparse vegetation. Adults and larvae are nocturnal, hiding in burrows during the day. They are strong burrowers and fast runners. Adults hibernate. References: Brett, pers comm., Erwin (2001), Benschoter & Cook (1956), Jeannel (1946) Larochelle & Larivière (2003), Lindroth (1961)

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Omophron americanum Dejean 1831 AMERICAN ROUND SAND BEETLE Omophron Americanus Dejean, 1831:583 Omophron Sayi Kirby, 1837:65 Omophron lacustrae Casey, 1897:301 Omophron texanus Casey, 1897:302 Omophron admiscens Casey, 1913:8 Omophron fontinale Casey, 1913:41 Omophron iridescens Casey, 1913:41 Omophron Lengi Casey, 1920:135 Omophron illustre Casey, 1920:136 Homophron tanneri Chandler, 1941:100

Distribution: Native, New World. CANADA – AB, MB, NB, NF, NS, ON, PE, PQ, SK; USA – AL, AR, AZ, CO, CT, DC, DE, FL, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NY, OH, OK, PA, RI, SC, SD, TN, TX, UT, VA, VT, WI, WV. Way of Life: M ACROHABITAT : Lowlands to uplands, 260 – 2195 meters altitude. MICROHABITAT: On open bare, or almost so, wet ground consisting of sand, or sand mixed with clay, sometimes saline, at the margins of rivers, brooks, lakes, ponds, pools, marshes, and estuaries. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in April – December; copulating pairs in April – May; gravid females in May – June; teneral adults in July – September. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground (adult and larva). Adults found over wintering on higher and drier ground, in the upper zone of river banks and sand pits, at the edge of woods and in the woods; in the ground, under dead leaves, stones, fallen trees, and bark. Adults are gregarious and often attracted to lights at night and they are found in drift material after flights over water. They also stridulate. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Omophron baenningeri Dupuis 1912 BÄNNINGER’S ROUND SAND BEETLE Omophron Bänningeri Dupuis, 1912:60

Distribution: Native, New World. MÉXICO – VC. Way of Life: MACROHABITAT: Midlands to uplands, 885 – 1981 meters altitude. MICROHABITAT: Stream margins. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March, July, and September – November. BEHAVIOR: Unknown. References: Erwin (2001), Data from CAS, UASM collections Omophron dentatum LeConte 1852 TOOTHED ROUND SAND BEETLE Omophron dentatus LeConte, 1852:200

Distribution: Native, New World. MÉXICO – BJ; USA – CA.

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Way of Life: MACROHABITAT: Lowlands to midlands, 30 – 1219 meters altitude. MICROHABITAT: On open wet, sandy or gravelly ground at the margin of rivers, brooks, and ponds. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in February – August, and October – November. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground (adult and larva). Adults occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Omophron dominicense Chaudoir 1868 INSULAR ROUND SAND BEETLE Omophron dominicense Chaudoir, 1868:58

Distribution: Native, New World. HISPANIOLA. Way of Life: Unknown. References: Erwin (2001), Roberta Brett, CAS (pers. comm.) Omophron gilae LeConte 1852 GILA RIVER ROUND SAND BEETLE Omophron gilae LeConte, 1852:201 Omophron pallidus Casey, 1897:305 Omophron pimalis Casey, 1913:44

Distribution: Native, New World. MÉXICO – SO; USA – AZ, CA, CO, NM, TX, UT. Way of Life: MACROHABITAT: Lowlands to midlands, 146 – 1372 meters altitude. MICROHABITAT: Margins of water bodies. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in January, and March – August. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground. Adults are gregarious. Adults occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Omophron gratum Chaudoir 1868 PLEASING ROUND SAND BEETLE Omophron gratus Chaudoir, 1868:58

Distribution: Native, New World. COSTA RICA; GUATEMALA; MÉXICO – CH, CS, JA, MH, NA, NL, OA, PU, SI, SL, SO, TM, VC. Way of Life: MACROHABITAT: Lowlands to uplands, 30 – 1524 meters altitude. MICROHABITAT: Along river margins. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – March, July – August, and November – December. BEHAVIOR: Unknown. References: Erwin (2001), Roberta Brett, CAS (pers. comm.), Data from CAS, NMNH, UASM collections

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Omophron grossum Casey 1909 LARGE ROUND SAND BEETLE Omophron grossus Casey, 1909:275

Distribution: Native, New World. USA – AR, IA, KS, MO, MS, OK, TX. Way of Life: MACROHABITAT: Midlands, 510 – 540 meters altitude. MICROHABITAT: On open wet, sandy or mixed with mud ground along river margins. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground, or in mole cricket burrows. Adults are gregarious and occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Omophron labiatum (Fabricius) 1801 LARGE-LIPPED ROUND SAND BEETLE Scolytus labiatus Fabricius, 1801:248

Distribution: Native, New World. CANADA – NS; USA – AL, CT, DC, FL, GA, LA, MA, MD, MS, NC, NJ, NY, PA, SC, TX, VA. Way of Life: MACROHABITAT: Coastal lowlands, sea level – 282 meters altitude. MICROHABITAT: On open wet, sandy, sometimes saline, bare or almost so, substrate in cultivated fields (e.g., corn), or margins of ponds, lakes, and rivers; margins of ditches; and sea beaches. DISPERSAL ABILITIES: Macropterous, adults occasionally found at lights at night. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Mostly nocturnal, although sometimes active in the sunshine; usually adults take cover during the day in burrows dug in the ground. Adults are gregarious. Larval food sometimes are grains and young shoots of corn, thus sometimes as a pest of young corn of economic importance. Adults occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), (Roberta Brett, CAS, pers. comm.) Data from CAS, NMNH collections Omophron mexicanum Dupuis 1912 MEXICAN ROUND SAND BEETLE Omophron mexicanus Dupuis, 1912:61

Distribution: Native, New World. MÉXICO – VC. Way of Life: Unknown. References: Erwin (2001), Roberta Brett, CAS (pers. comm.) Omophron nitidum LeConte 1848 SHINY ROUND SAND BEETLE Omophron nitidus LeConte, 1848:347 Omophron nitens Chaudoir, 1868:60

Distribution: Native, New World. USA – AL, AR, GA, IA, IL, IN, KS, LA, MI, MN, MO, MS, NE, OK, TN, TX.

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Way of Life: MACROHABITAT: Lowlands, 10 – 260 meters altitude, on beaches of lakes and rivers. MICROHABITAT: On open wet, muddy ground, bare or covered with sparse vegetation. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in March – December. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground, or in leaf litter. Adults are gregarious and often attracted to lights. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Omophron obliteratum G. Horn 1870 EFFACED ROUND SAND BEETLE Omophron obliteratus G. Horn, 1870:73 Omophron sonorae Casey, 1897:304 Omophron utense Casey, 1913:43 Omophron subimpressus Casey, 1913:43

Distribution: Native, New World. MÉXICO – CH, DU, SO, ZA; USA – AZ, CA, MT, NM, TX, UT. Way of Life: MACROHABITAT: Lowlands to mountains, 457 – 2256 meters altitude, in oak pine forest along creeks in canyons. MICROHABITAT: Open ground on creek and river margins. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in April – November. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground. Adults are gregarious. Adults repeatedly found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Omophron oblongiusculum Chevrolat 1835 LONG ROUND SAND BEETLE Omophron oblongiusculus Chevrolat, 1835: no. 166

Distribution: Native, New World. MÉXICO – CS, OA. Way of Life: MACROHABITAT: Midlands to uplands, 640 – 2000 meters altitude. MICROHABITAT: On sandy stream margins. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – March, June – August, and November – December. BEHAVIOR: Adults take cover in flood debris of Alnus-Salix along stream. References: Erwin (2001), Roberta Brett, CAS (pers. comm.), Data from CAS, NMNH, UASM collections Omophron ovale G. Horn 1870 OVAL ROUND SAND BEETLE Omophron ovale G. Horn, 1870:75 Omophron concinnum Casey, 1897:302 Omophron gemma Casey, 1897:304 Omophron frater Casey, 1913:41

Distribution: Native, New World. CANADA – AB, BC, SK; USA – CA, ID, MT, OR, UT, WA, WY. Way of Life: MACROHABITAT: Lowlands to uplands, 61 – 1509 meters altitude. MICROHABITAT: On open wet sandy bare ground, often mixed with clay along margins of rivers, brooks, lakes,

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ponds, and reservoirs. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in April – October; copulating pairs in July; teneral adults in July and October. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground, or under stones, or vegetal debris. Adults are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Omophron robustum G. Horn 1870 ROBUST ROUND SAND BEETLE Omophron robustus G. Horn, 1870:73 Omophron brevipenne Casey, 1909:276 Omophron decoloratus Fall, 1920:211

Distribution: Native, New World. CANADA – AB, ON; USA – IA, IL, IN, KS, MI, MN, ND, NE, OH, OK, SD, WI . Way of Life: MACROHABITAT: Lowlands, 57 – 670 meters altitude, on beaches of lakes, rivers, and brooks. MICROHABITAT: On open wet, sandy, sometimes alkaline ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in August. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground or under debris. Adults are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Omophron solidum Casey 1897 SOLID ROUND SAND BEETLE Omophron solidus Casey, 1897:303 Omophron lawrencei Hatch, 1953:69

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, 119 – 1052 meters altitude, beaches of water bodies. MICROHABITAT: Moist sandy ground near water. D ISPERSAL ABILITIES : Macropterous, probably capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in April – November. BEHAVIOR: Nocturnal, adults take cover during the day in burrows dug in the ground. Adults are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Omophron sphaericum Chevrolat 1835 ROUND ROUND SAND BEETLE Omophron sphaericus Chevrolat 1835: no. 167

Distribution: Native, New World. GUATEMALA; MÉXICO – VC . Way of Life: Unknown. References: Erwin (2001), Roberta Brett, CAS (pers. comm.)

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Omophron tessellatum Say 1823 MOSAIC ROUND SAND BEETLE Omophron tessellatus Say, 1823:152 Omophron Lecontei Dejean, 1831:582 Omophron tesselatus LeConte, 1848:447 Omophron ellipticus Casey, 1909:276

Distribution: Native, New World. CANADA – AB, MB, NB, NS, ON, PE, PQ, SK; USA – AZ, CA, CO, CT, IA, IL, IN, KS, KY, MA, ME, MI, MN, MO, MT, ND, NE, NH, NJ, NM, NY, OH, PA, RI, SD, TX, VA, VT, WI, WY. Way of Life: MACROHABITAT: Lowlands to uplands, 79 – 1801 meters altitude. MICROHABITAT: On open wet, sandy or clayish, sometimes saline ground, bare or almost so, on margins of lakes, ponds, estuaries, slow brooks, and rivers; sea beaches; margins of pools (e.g., in sand pits) and ditches. DISPERSAL ABILITIES: Macropterous, capable of flight. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in April – November; copulating pairs in June – July; teneral adults in August – October. BEHAVIOR: Mostly nocturnal, sometimes active in the sunshine, usually adults take cover during the day in burrows dug in the ground (adult and larva). Adults are gregarious and often attracted to lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections LORICERITAE Loricerini LeConte 1861 SPRINGTAIL HUNTERS Loricerini LeConte, 1861:6

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Elaphrini Latreille 1802 Geographic status: Holarctic, Neotropical. References: Ball & Erwin (1969), Erwin (2001) Loricera Latreille 1802 SPRINGTAIL HUNTERS (Plate 12) Loricera Latreille, 1802:88 Lorocera Agassiz, 1847:216

Number of species: 13. Number of Western Hemisphere species: Five. Taxonomy: Stable. Adelphotaxon: See Loricerini, above. Type species: Carabus pilicornis, Fabricius, 1775. Distribution: Native, New World (four species): Holarctic (one species), Nearctic (four species), Neotropical (one species), Palearctic (eight species).

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Habitat: According to Lindroth (1961), adults of this genus are very hygrophilous and very active in the sunshine, running and flying. Adults are predaceous on springtails (Collembola), as are the larvae that possess adhesive mouthparts. Adults use their stiff long setae on the basal segments of their antennae as an enclosing trap to capture springtails. Larvae use the apical part of their galea as a sticky trap to capture springtails. Adults hibernate. References: Ball & Erwin (1969), Erwin (2001), Lindroth (1961) Loricera aptena Ball & Erwin 1969 WINGLESS SPRINGTAIL-HUNTER Loricera aptena Ball & Erwin, 1969:889

Distribution: Native, New World. MÉXICO – CH, DU, MH, SI. Way of Life: MACROHABITAT: Uplands to mountains, 2150 – 2850 meters altitude, in oak/ pine/fir forests, pine/oak forests with Alder, Madrone and Juniper, and coniferous forests. MICROHABITAT: Adults are found in leaf litter, or under fallen trees and stones. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running, not capable of flight. Good runners. SEASONAL OCCURRENCE: Adults have been found in July – November. A larva and three pupae were also found in September. BEHAVIOR: See description under genus, above. Adults take cover in the day under fallen trees and in leaf litter. References: Ball & Erwin (1969), Erwin (1991, 2001), Data from CAS, NMNH, UASM collections Loricera decempunctata Eschscholtz 1833 10-SPOTTED SPRINGTAIL-HUNTER Loricera decempunctata Eschscholtz, 1833:25

Distribution: Native, New World. CANADA – BC, QC; USA – AK, CA, OR, WA. Way of Life: MACROHABITAT: Lowlands to midlands, subalpine, and alpine zones, 2 – 585 meters altitude. MICROHABITAT: Adults of this species occur near fresh water (e.g. borders of ponds, lakes, marshes, slow rivers and brooks, and seepages), but they are not as hygrophilous as L. pilicornis adults; they also are often found among grass and moss in the open some distance from the waters edge, in cultivated fields and gardens. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in February, and May – October. BEHAVIOR: See description under genus, above. Adults are mostly nocturnal, but also run in the sunshine, and occasionally fly. Adults hibernate. References: Erwin (2001), Kavanaugh (1992), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Loricera foveata LeConte 1851 PITTED SPRINGTAIL-HUNTER Loricera foveata LeConte, 1851:180

Distribution: Native, New World. CANADA – BC; USA – CA, OR, WA. Way of Life: MACROHABITAT: Lowlands to midlands, 20 – 853 meters altitude. MICROHABITAT: Adults of this species occur in the vicinity of fresh water (e.g. borders of reservoirs), but they

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are not as hygrophilous as L. pilicornis adults; they are often under stones, boards, and fallen trees on clay-mixed sandy ground up slope from the water’s edge; clay ground at edges of vernal pools and Juncus marshes. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January, March – August, and November – December. BEHAVIOR: See description under genus, above. Adults are mostly nocturnal, but also run in the sunshine. Adults are predaceous. References: Erwin (2001), Erwin (collecting notes), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Loricera pilicornis congesta von Mannerheim 1853 BERING SPRINGTAIL-HUNTER Loricera pilicornis congesta von Mannerheim, 1853:121

Distribution: PALEARCTIC; USA – AK. Way of Life: MACROHABITAT: Coastal lowlands and islands, sea level – 29 m altitude. MICROHABITAT: Pronouncedly hygrophilous, at the border of fresh standing water with partially shaded rich organic ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Loricera pilicornis pilicornis (Fabricius) 1775 HAIRY-HORNED SPRINGTAIL-HUNTER Carabus pilicornis Fabricius, 1775:243 Carabus seticornis O.F. Müller, 1776:79 Loricera vinearum Geoffroy, 1785:43 Loricera aenea Latreille, 1806:224 Loricera semipunctata Eschscholtz, 1833:25 Loricera alpina Heer, 1838:44 Loricera rufilabris Motschulsky, 1845:340 Loricera californica LeConte, 1863:3 Loricera neoscotica LeConte, 1863:3 Loricera uteana Casey, 1920:147 Loricera sierrae Van Dyke, 1925:113 Loricera apennina Binaghi, 1942:135 Loricera semistriata Hatch, 1953:64

Distribution: Palearctic; Canada – AB, BC, LB, MB, NB, NF, NS, NT, ON, PE, PM, PQ, SK, YT; USA – AK, CA, CO, ID, IN, MA, ME, MI, MN, MT, ND, NE, NH, NJ, NM, NY, OH, OR, PA, SD, UT, VT, WA, WI, WY. Way of Life: MACROHABITAT: Lowlands to mountains, 274 – 2743 meters altitude. MICROHABITAT: Hygrophilous and eurytopic, on soft, wet, often muddy ground rich in organic debris, covered with sparse vegetation and shaded by shrubs and tall plants at the borders of eutrophic marshes, ponds, lakes, pools (often polluted by refuse), slow rivers, brooks, and

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irrigation ditches; fens, swamps, Carex marshes, low forests, flood-plain forests, mud flats, wet meadows and pastures, cultivated fields (e.g., barley, dry bean). Also, found in apple orchards in BC and gardens in CA, thus not dependent on water. Moderately favored by human activities. DISPERSAL ABILITIES: Macropterous, adults occasionally found at lights at night. Frequent in lake-and seashore drift material, Occasional climber on plants and trees. SEASONAL OCCURRENCE: Adults have been found in January – December; copulating pairs in May and July; teneral adults in June – September. BEHAVIOR: Diurnal and nocturnal, active in the sunshine or adults take cover during the day at the base of plants, under plant mats, heaps of grass, dead leaves, stones, pieces of wood, and in moss. Adults are gregarious in winter. Adults produce acoustic signals. Adults found over wintering on higher and drier ground, in the upper banks of marshes and swamps, in woods, at the edge of woods, in vacant lots, and on hillocks; under dead leaves, stones, fallen trees, fallen pieces of bark, under the loose bark of fallen trees, in rotten standing tree stumps, and in the ground (as deep as 20 cm). Adults are predaceous on springtails and insect larvae. Larvae are also predaceous on springtails, and mites. References: Erwin (2001), Forsythe (1979), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections Loricera rotundicollis Chaudoir 1863 ROUND-COLLARED SPRINGTAIL-HUNTER Loricera rotundicollis Chaudoir, 1863:115

Distribution: Native, New World. GUATEMALA; MÉXICO – GO, JA, HD, MX, MH, ML, OA, PU, QT, SL, SI, VC. Way of Life: M ACROHABITAT : Midlands to mountains, 930 – 2928 meters altitude. MICROHABITAT: Adults are found on higher, dry ground in cloud, oak, oak-pine forests with Arbutus, Arctostaphylos, Abies, Quercus spp. and Pinus ponderosa Douglas ex Lawson & C. Lawson, P. durangensis Martínez, P. cooperi C.E. Blanco; rarely found in open areas at higher altitudes. DISPERSAL ABILITIES: Wing-dimorphic, brachypterous the most common form; macropterous individuals found only in large populations. SEASONAL OCCURRENCE: Adults have been found in January, March, and May – December. Larvae found in August. BEHAVIOR: See description under genus, above. Adults take cover in the day under fallen trees and in the leaf litter. Larvae and pupae also found under large embedded stones. References: Ball & Erwin (1969), Erwin (2001), Data from CAS, NMNH, UASM collections CARABITAE Carabini Latreille 1802 SEARCHERS Carabini Latreille, 1802:80

Number of worldwide genera: Eight. Number of Western Hemisphere genera: Five. Taxonomy: Unstable, many new species yet to be described; no global synthesis. Adelphotaxon: Cychrini Laporte de Castelnau 1834 Geographic status: All Realms. References: Erwin (2001), Lorenz (2005)

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Carabina Latreille 1802 SEARCHERS Carabini Latreille, 1802:80

Number of worldwide genera: Seven. Number of Western Hemisphere genera: Five. Taxonomy: Unstable in part, many new species yet to be described in the Palearctic; no global synthesis. Adelphotaxon: Ceroglossina Vacher de Lapouge 1927 Geographic status: Holarctic, Neotropical, Neaustral, Oriental, Australasian, Afrotropical References: Erwin (2001), Lorenz (2005) Notes on the Calasoma-group: Gidaspow (1959) summarizes the feeding habits of the Calasoma-generic group of species in the Western Hemisphere, as follows: “The larvae of Calosoma are very active when the weather is hot and feed by day and night. They usually attack the caterpillars from beneath or from the side, but if the caterpillars are hairy, they pinch them between the segments. Even a newly hatched larva is able to kill a caterpillar of considerable size. The larvae eat the liquid and fatty parts of the caterpillars and often leave the meal before it is finished, thus destroying more caterpillars than they actually need. The larvae favor the pupae of butterflies, except those that are hidden in cocoons. However, there are records that cocoons of Malacosoma americana and Malacosoma disstria have been destroyed. Usually a larva makes a hole on the side of the pupa, between the segments, enlarges it, and then pushes the front part of its body inside. It leaves the pupa when there is virtually nothing left but a shell. While eating, the larva is well hidden beneath the old pupa cases. The entrance hole made by the larva is always irregular in outline and cannot be confused with holes made by parasites. Preference is given to the pupae of females. There are records (Burgess, 1911) stating that among the pupae killed by the larvae of Calosoma sycophanta 75 per cent were females. One larva of Calosoma as large as sycophanta or scrutator is able to kill about 50 caterpillars or 15 pupae between the time of hatching and pupation. The adults of Calosoma seize the caterpillars in the middle of the back and easily kill them in a few seconds regardless of size. During the season, one beetle can destroy about 300 caterpillars. Actually, some species, such as Calosoma sycophanta, which live from two to three years, destroy even more caterpillars. If another beetle is attacked, it is often seized by the neck between the occiput and pronotum, or, if it is a beetle with a soft abdomen, such as some species of the family Chrysomelidae, it is attacked from below. In captivity, the beetles can also eat beef meat, but after a week or so, they refuse to feed, and caterpillars must be given to them. Apparently, the species of Calosoma are not affected by the diseases from which some caterpillars often suffer. Experiments showed that Calosoma sycophanta was not affected by a disease of the gypsy moth known as “wilt.” Cannibalism among the larvae is not uncommon and, to a certain extent, reduces the number of larvae even when food is abundant. The best known species of the genus, Calosoma sycophanta, is, as is said above, more or less adapted to the life of the gypsy moth and is very active when the caterpillars of the gypsy moth are present. Most of the larvae hatch during the pupal stage of the moth. In spite of the fact that sycophanta attacks other caterpillars and pupae, it is never numerous in the regions where the gypsy moth is absent. Both larvae and adults are very good climbers; they mass on trunks of the trees under branches where the pupae of the gypsy moth are usually hidden and devour them on the spot. Occasionally they attack the imago also, mostly a female laying eggs. Calosoma frigidum lives chiefly on Heterocampa guttivitta and other Lepidoptera that pupate on the ground. The larvae of frigidum are not good climbers and feed on pupae that are usually hidden in cells under leaf mold. Calosoma calidum is also more terrestrial in habits; the

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larvae do not often climb but usually prey on caterpillars and pupae that stay on the ground. The adults, however, climb trees easily. Records show that they destroy cankerworms, cutworms, army worms, locusts, and Colorado potato beetles. The larvae of Calosoma cancellatum are also adapted to those insects that stay on the ground. Calosoma scrutator and haydeni, on the contrary, feed on various caterpillars that live on trees and bushes, although there are indications that they are not such good climbers as is Calosoma sycophanta. Calosoma externum is known to feed on locusts and army worms.” Callisthenes Fischer von Waldheim 1820 BEAUTIFUL BLACK SEARCHERS (Plate 13) Callisthenes Fischer von Waldheim, 1820: pl. 7 Chrysostigma Kirby, 1837:19 [valid subgenus for native Nearctic and Neotropical species] Tapinosthenes Kolbe, 1895:56 Microcallisthenes Apfelbeck, 1918:161 Callistenia Vacher de Lapouge, 1929:2 [valid subgenus for native Nearctic species] Lyperostenia Vacher de Lapouge, 1929:3 See Lorenz (2005) for additional Old World subgenera.

Number of worldwide species: 53. Number of Western Hemisphere species: 25. Taxonomy: Stable. Adelphotaxon: Calosoma Weber 1801 Type species: Callisthenes panderi Fischer von Waldheim, 1820 Distribution: Holarctic. Native, New World (24 species), Palearctic (29 species) Habitat: According to Lindroth (1961), most species of this genus occur in open dry country. Adults of subgenus Callistenia have reduced wings and are flightless, those of Chrysostigma are fully winged and fly, the latter also are climbers. The life history of many species in North America is known (Burgess & Collins, 1917). Adults and larvae are ravenous carnivores, specializing upon caterpillars and pupae of Lepidoptera, never, as far as is known, monophagously restricted to a single host. References: Erwin (2001), Gidaspow (1959), Lindroth (1961) Callisthenes (Chrysostigma) affinis (Chaudoir) 1843 RELATED BEAUTIFUL BLACK SEARCHER Calosoma affine Chaudoir, 1843:746 Calosoma triste LeConte, 1845:201 Calosoma tristoides Fall, 1910:92

Distribution: Native, New World. MÉXICO – CH, DU, DF, HD, MX, NL, OA, SL, TM; USA – AR, AZ, CA, CO, KS, MO, MN, NE, NM, NV, OK, TX, UT. Way of Life: MACROHABITAT: Lowlands to mountains, 9 – 2417 meters altitude. MICROHABITAT: Adults are found in pastures, cultivated fields (e.g. alfalfa), and sand dunes, on open sandy ground with sparse vegetation. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Moderate runners, and climbers on plants. SEASONAL OCCURRENCE: Adults have been found in April – October. Females with eggs in July – August. BEHAVIOR: Adults are gregarious (sometimes

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swarming) and are predaceous on lepidopterous caterpillars. They are both diurnal and nocturnal, adults take cover in the day in the ground and under debris. Larvae hibernate. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Callisthenes (Chrysostigma) ampliator (Bates) 1891 LARGE BEAUTIFUL BLACK SEARCHER Calosoma ampliator Bates, 1891:223

Distribution: Native, New World. MÉXICO – CH, DU, GJ, SO, TM, VC. Way of Life: MACROHABITAT: Uplands, 1,100 – 1420 meters altitude. MICROHABITAT: Likely near wet or damp areas with shrubby vegetation or small trees in the Chihuahuan Desert and possibly at the margins of cultivated lands. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (1991, 2001), Gidaspow (1959) Notes. The record from Panamá in Gidaspow (1959) is doubtful (see Erwin, 1991). Callisthenes (Chrysostigma) calidus (Fabricius) 1775 FIERY HUNTER Carabus calidus Fabricius, 1775:237 Calosoma calida expansa Casey, 1897:344 Calosoma calida laticollis Casey, 1897:344 Calosoma calida stellata Casey, 1897:344 Calosoma comes Casey, 1920:156 Calosoma concreta Casey, 1920:157 Chrysostigma lepidum ocellatum Vacher de Lapouge, 1931:381

Distribution: Native, New World. CANADA – AB, BC, MB, NB, NS, NT, ON, PM, PQ, SK; PUERTO RICO; USA – AL, AR, CO, CT, DC, DE, GA, IA, IL, IN, KS, KY, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NM, NY, OH, OR, PA, RI, SD, TN, UT, VA, VT, WA, WI, WV, WY Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 700 meters altitude. MICROHABITAT: Adults of this species are ground-dwelling, often in leaf litter at base of trees and inhabitants of open rather dry fields with low vegetation, and cultivated lands in many crops. DISPERSAL ABILITIES: They are fully winged and probably fly quite well; they are attracted to lights. SEASONAL OCCURRENCE: Adults have been found in March to December. Adults over winter and are found in the late spring and throughout the summer (seen from May). Larvae are present in the fauna in summer. BEHAVIOR: Adults may live for up to 3 years, but this is not common. These are poor climbers, but will go up the trunk of trees a short ways for caterpillars (e.g. Porthetria dispar, Hyphantria cunea), occasionally but not in the larval stage. Both adults and larvae are predaceous on cutworms, and they will also eat tent caterpillars and army worms (Burgess and Collins, 1917). They can be collected by tying burlap sacks around the lower trunks of trees, a device that congregates their prey; they will also come to sugar used for “sugaring” for moths. They are both nocturnal, and according to Larochelle (1975), diurnal, as well. Only one specimen was collected on Plummers Island in Maryland, in 1913 “at light.” There have been no recent records. Hibernation takes place in the soil up to 6 cm deep. A long and detailed report on their bionomics is given by Burgess and Collins (1917) and is briefly summarized in Larochelle & Larivière (2003).

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References: Erwin (2001), Burgess and Collins (1917), Larochelle (1975), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Callisthenes (Chrysostigma) cancellatus (Eschscholtz) 1833 LATTICED BEAUTIFUL BLACK SEARCHER Calosoma cancellatum Eschscholtz, 1833:23 Calosoma aenescens LeConte, 1854:16 Calosoma esuriens Casey, 1913:64 Calosoma transversa Casey, 1913:65 Calosoma sagax Casey, 1920:158 Calosoma rectilatera Casey, 1920:158 Calosoma praestans Casey, 1920:159

Distribution: Native, New World. CANADA – BC; USA – AZ, CA, ID, MT, NV, OR, UT, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 3 – 1371 meters altitude. MICROHABITAT: On open dry ground, often adults are found in cultivated fields (e.g. sugar beet), also in tule marshes in CA. DISPERSAL ABILITIES: Adults are submacropterous, probably incapable of flight and are moderate runners and occasional climbers (on shrubs). SEASONAL OCCURRENCE: Adults are active in March – September; females with eggs in May – June. Larvae are present in the fauna in summer and thus over winter. BEHAVIOR: Adults are diurnal; active in the sunshine on the ground, or on low shrubbery. Adult food items, in the field, are lepidopterous caterpillars, elaterids, flies, and orthopterans. The larva lives on, or near the ground and is known to feed upon larvae of the moth Peridroma saucia (Hübner). References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Callisthenes (Callistenia) dawsoni Dajoz 1997 DAWSON’S BEAUTIFUL BLACK SEARCHER Callisthenes dawsoni Dajoz, 1997:70

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Big Alkali Lake, 2100 meters altitude. MICROHABITAT: On open ground which is covered almost everywhere with calcium salts and with sparse vegetation cover dominated by Graminea and some Artemisia shrubs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in August. BEHAVIOR: Adults are predaceous on caterpllars. References: Dajoz (1997), Erwin (2001) Callisthenes (Callistenia) dietzi (Schaeffer) 1904 DIETZ’S BEAUTIFUL BLACK SEARCHER Calosoma Dietzii Schaeffer, 1904:197 Callisthenes gravidulus Casey, 1913:68

Distribution: Native, New World. USA – CA.

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Way of Life: M ACROHABITAT : Midlands to mountains, 800 – 2700 meters altitude. MICROHABITAT : Open dry grasslands and mountain meadows. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – July. BEHAVIOR: Adults climb grasses. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Callisthenes (Callistenia) discors (LeConte) 1857 DIFFERENT BEAUTIFUL BLACK SEARCHER Calosoma discors LeConte, 1857:31 Callisthenes discors inversus Casey, 1913:67

Distribution: Native, New World. USA – CA, WA. Way of Life: MACROHABITAT: Lowlands to midlands, 100 – 1200 meters altitude, on coastal grasslands and in the Sierra Nevada foothills in oak savannahs. MICROHABITAT: On the ground amongst grasses. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – July. BEHAVIOR: Unknown. References: Erwin (2001), Jeannel (1940), Larochelle & Larivière (2003), Data from NMNH collection Callisthenes (Callistenia) lariversi (Van Dyke) 1943 LARIVERS’ BEAUTIFUL BLACK SEARCHER Calosoma lariversi Van Dyke, 1943:17

Distribution: Native, New World. USA – CA, NV. Way of Life: MACROHABITAT: Uplands, 1750 meters altitude, in the Great Basin desert. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Callisthenes (Callistenia) latipennis (G. Horn) 1870 BROAD BEAUTIFUL BLACK SEARCHER Calosoma latipenne G. Horn, 1870:70 Calosoma arcuata Casey, 1897:343 Callisthenes tularensis Casey, 1913:68 Callisthenes optimus Casey, 1913:69

Distribution: Native, New World. USA – CA, NV. Way of Life: MACROHABITAT: Lowlands to midlands, 113 – 1300 meters altitude, in Mojave Desert vegetation. MICROHABITAT: Sandy soils with desert scrub in lowlands; sandy soils and rocky terrain in Juniper-Pinyon pine open forests on the foothills. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults are found in January – July. BEHAVIOR: Adults feed on grasshoppers. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

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Callisthenes (Chrysostigma) lepidus (LeConte) 1845 NEAT BEAUTIFUL BLACK SEARCHER Calosoma lepidum LeConte, 1845:201

Distribution: Native, New World. CANADA – AB, MB, SK; USA – MT, ND, SD, WY. Way of Life: MACROHABITAT: Midlands, 1300 meters altitude, in short grass open prairies. MICROHABITAT: Open ground; rather dry ground covered with some vegetation, such as grasses, and in cultivated fields and along their borders (e.g. wheat). DISPERSAL ABILITIES: Subbrachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Callisthenes (Callistenia) luxatus (Say) 1823 DISLOCATED BEAUTIFUL BLACK SEARCHER Calosoma luxata Say, 1823:149 Carabus Zimmermanni LeConte, 1848:345 Calosoma striatulum LeConte, 1859:4 Calosoma pimelioides Walker, 1866:312 Callisthenes luxatus opacus Géhin, 1885:70 Callisthenes exaratus Casey, 1913:72 Callisthenes tegulatus Casey, 1913:72 Callisthenes tegulatus viator Casey, 1913:72 Callisthenes pustulosus Casey, 1913:73 Callisthenes diffractus Casey, 1913:75 Callisthenes reflexus Casey, 1920:164 Callisthenes utensis Casey, 1920:165 Callisthenes semotus Casey, 1920:166 Callisthenes debilis Casey, 1920:167 Callisthenes parowanus Casey, 1920:167 Calosoma luxatum zimmermanni striatus Breuning 1928:86 Calosoma zimmermanni tahoensis Van Dyke, 1943:18

Distribution: Native, New World. CANADA – AB, BC, SK; USA – AZ, CA, CO, ID, KS, MT, NE, NM, NV, OR, SD, UT, WA, WY. Way of Life: MACROHABITAT: Midlands to mountains, alpine zone, 823 – 4000 meters altitude. MICROHABITAT: On open short grass praire, and on open ground with sparse vegetation within dry forests (e.g., Pinus). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – September. BEHAVIOR: Adults take cover under stones. References: Dajoz (1997), Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from CAS, NMNH collections

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Callisthenes (Callistenia) moniliatus LeConte 1852 COLLARED BEAUTIFUL BLACK SEARCHER Callisthenes moniliatus LeConte, 1852:200 Calosoma laqueatum LeConte, 1860:318 Carabus bicolor Walker, 1866:313 Callisthenes concinnus Casey, 1913:66 Carabus taedatus vancouvericus Csiki, 1927:286

Distribution: Native, New World. CANADA – AB, BC, SK; USA – AZ, CA, ID, MT, NE, OR, WA. Way of Life: MACROHABITAT: Uplands, 1800 – 2200 meters altitude. MICROHABITAT: On open ground with sparse vegetation on short grass praire and in dry forests (e.g., Pinus). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – August. BEHAVIOR: Adults take cover under stones. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Callisthenes (Callistenia) monticola (Casey) 1897 MOUNTAIN BEAUTIFUL BLACK SEARCHER Calosoma monticola Casey, 1897:342 Callisthenes nevadensis Casey, 1913:74

Distribution: Native, New World. USA – CA, ID, NV, OR, UT, WA, WY. Way of Life: MACROHABITAT: Midlands, 1067 – 1219 meters altitude. MICROHABITAT: Open dry ground in sagebrush areas (Artemisia sp.). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – June, and August – September. BEHAVIOR: Adults are predaceous on cutworms, wireworms, scarabaeids, coccinelids, meloids, and staphylinds. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Callisthenes (Chrysostigma) morrisonii (G. Horn) 1885 MORRISON’S BEAUTIFUL BLACK SEARCHER Calosoma Morrisonii G. Horn, 1885:128 Calosoma mexicanum Géhin, 1885:67

Distribution: Native, New World. MÉXICO – DU; USA – CA, CO, NM, NV. Way of Life: MACROHABITAT: Uplands, 1520 – 1857 meters altitude. MICROHABITAT: Open dry ground, sand dunes. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – May. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

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Callisthenes (Chrysostigma) obsoletus (Say) 1823 OLD BEAUTIFUL BLACK SEARCHER Calosoma obsoleta Say, 1823:149 Calosoma indistinctum LeConte, 1845:208 Calosoma obsoleta microsticta Casey, 1897:345

Distribution: Native, New World. CANADA – AB, MB, SK; USA – CO, IA, ID, KS, MN, MT, ND, NE, NM, NV, OR, SD, UT, WY Way of Life: MACROHABITAT: Lowlands to uplands, 321 – 2186 meters altitude. MICROHABITAT: On open ground covered with some vegetation (e.g., grass or weeds) on open prairies, rangelands, cultivated fields (e.g., alfalfa, corn, wheat), along fence rows, and roadsides. Markeldy favored by human activities. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – November; oviposition in June – August. BEHAVIOR: Crepuscular, adults take cover during the day under pieces of wood, dry cow dung and dead prostrate weeds; sometimes active by the hundreds on cloudy days. Adults found overwintering in cavities in the ground (as deep as 15 cm). Life span: probably 2 – 3 years. Adults and larvae are predaceous on lepidopterous caterpillars and under lab conditions will eat grasshoppers. The favored prey of adults are the larvae of the moth, Hemileuca oliviae Ckll; that of the larvae, the cocooned pupa of this moth species. References: Erwin (2001), Burgess & Collins (1917), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Callisthenes (Callistenia) oregonus Gidaspow 1959 OREGON BEAUTIFUL BLACK SEARCHER Callisthenes oregonus Gidaspow, 1959:317

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959) Callisthenes (Callistenia) placerus Gidaspow 1959 PLACER BEAUTIFUL BLACK SEARCHER Callisthenes placerus Gidaspow, 1959:309

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Uplands, 1802 – 2134 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

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Callisthenes (Callistenia) schaefferi (Breuning) 1928 SCHAEFFER’S BEAUTIFUL BLACK SEARCHER Calosoma irregulare Schaeffer, 1915:235 Calosoma discors schaefferi Breuning, 1928:79 Calosoma striatius Hatch, 1953:54, NEW SYNONYMY

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Midlands to uplands, 600 – 1981 meters altitude. MICROHABITAT: On the ground in damp meadows. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Unknown. References: Erwin (2001), Data from CAS, NMNH collections Note: Gidaspow (1959) states that she did not see Hatch’s type, nor could she make much of the description. I have compared the Holotype of C. striatus (now in NMNH) with Scheaffer’s type of C. irregulare (also in NMNH) and they represent the same species. Callisthenes (Chrysostigma) semilaevis (LeConte) 1852 SEMI-SMOOTH BEAUTIFUL BLACK SEARCHER Calosoma semilaeve LeConte, 1852:199 Calosoma semilaeve davidsoni Casey, 1914:33 Calosoma semilaeve adjutor Casey, 1920:162

Distribution: Native, New World. MÉXICO – BJ (+ GUADALUPE ISLAND); USA – AZ, CA, ID, OR, UT. Way of Life: MACROHABITAT: Lowlands to midlands, 4 – 1342 meters altitude. MICROHABITAT: Cultivated fields (e.g., sugar beet, clover), road sides, ground under low shrubs, and among peavines. Markedly favored by human activities. DISPERSAL ABILITIES: Macropterous, probably capable of flight. Adults and larvae are occasional climbers. SEASONAL OCCURRENCE: Adults have been found in April – August; copulating pairs in April; gravid females in May – June. BEHAVIOR: Diurnal and nocturnal. Adults found overwintering in cavities in the ground (7.5 – 10 cm deep). Adult are predaceous on caterpillars (e.g. Peridroma saucia Hübner and Malacosoma incurvum Hy Edwards), tenebrionids, elaterids, and carabids. Adults are most active in April – May. References: Erwin (2001), Burgess & Collins (1917), Larochelle & Larivière (2003), Data from NMNH collection Callisthenes (Chrysostigma) simplex (LeConte) 1878 SIMPLE BEAUTIFUL BLACK SEARCHER Calosoma simplex LeConte, 1878:61

Distribution: Native, New World. MÉXICO – ; USA – AZ, CA, CO, NM, TX. Way of Life: MACROHABITAT: Lowlands to uplands, 16 – 2195 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – May, August, and October – November. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Notes: Burgess & Collins (1917) reported this species from AZ, CA, CO, NM, TX, and MÉXICO, however, Gidaspow (1959) believes the species to be localized in central California.

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Callisthenes (Callistenia) subaeneus (Chaudoir) 1869 COPPERY BEAUTIFUL BLACK SEARCHER Calosoma subaeneum Chaudoir, 1869:28

Distribution: Native, New World. CANADA – BC; USA – CA, ID, WA. Way of Life: MACROHABITAT: Lowlands, 102 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – March. BEHAVIOR: Unknown. References: Erwin (2001), Data from CAS collection Callisthenes (Callistenia) subasperatus (Schaeffer) 1915 ROUGH BEAUTIFUL BLACK SEARCHER Calosoma subasperatum Schaeffer, 1915:235 Callisthenes klamathensis Casey, 1920:169

Distribution: Native, New World. USA – CA, NV, OR. Way of Life: MACROHABITAT: Midlands, 1248 – 1345 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – May. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Callisthenes (Chrysostigma) tepidus (LeConte) 1852 LUKEWARM BEAUTIFUL BLACK SEARCHER Calosoma tepidum LeConte, 1852:199 Calosoma irregulare Walker, 1866:312 Calosoma tepida caelator Casey, 1913:61 Calosoma tepida indigens Casey, 1913:61 Calosoma pellax Casey, 1920:160 Calosoma semicuprea Casey, 1920:161 Calosoma cogitans Casey, 1920:161

Distribution: Native, New World. CANADA – BC; USA – AZ, CA, CO, ID, MT, ND, NE, NV, OR, UT, WA, WY. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1524 meters altitude, sometimes near the sea. MICROHABITAT: On open dry ground covered with sparse low vegetation in cultivated fields (e.g, pea), rangelands, and along roadsides. DISPERSAL ABILITIES : Submacropterous, probably incapable of flight. SEASONAL OCCURRENCE : Adults have been found in May – August; teneral adults in May. BEHAVIOR: Adult are predaceous on caterpillars and grasshoppers. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection

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Callisthenes (Callistenia) wilkesii LeConte 1852 WILKES’S BEAUTIFUL BLACK SEARCHER Callisthenes Wilkesii LeConte, 1852:200

Distribution: Native, New World. CANADA – BC; USA – CA, ID, OR, WA. Way of Life: MACROHABITAT: Midlands, 732 – 900 meters altitude, in fields and meadows sometimes surrounded by woodlands. MICROHABITAT: Adults are found on dry ground in the open with low vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – September. BEHAVIOR: Nocturnal, adults take cover under under fallen trees in the daytime. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections Calopachys Haury 1880 BEAUTIFUL THICKISH SEARCHERS (Plate 15) Calopachys Haury, 1880:164 Eutelodontus Géhin, 1881: cxxxii

Number of worldwide species: 4. Number of Western Hemisphere species: 4. Taxonomy: Stable. Adelphotaxon: Calosoma Weber 1801 + ? Type species: Calopachys viridissimus Haury, 1880 Distribution: Neotropical (4 species). Although far enough south to be Neotropical, all of the species live high enough in altitude to be classified as dwelling in the temperate or subtropical zones. Habitat: Upland oak-pine forests of the Transverse Volcanic Belt of southern México. References: Erwin (2001), Gidaspow (1959), Korell & Frisch (2004) Calopachys blaptoides (Putzeys) 1846 HURTFUL BEAUTIFUL THICKISH SEARCHER Calosoma blaptoides Putzeys, 1846:400 Eutelodontus aztecus Géhin, 1881:cxxxii Eutelodontus depressicolle tehuacanus Vacher de Lapouge, 1924:40

Distribution: Native, New World. MÉXICO – GJ, OA, PU, VC. Way of Life: MACROHABITAT: Uplands to mountains, 2134 – 3320 meters altitude, in oak-pine forests and meadows. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal, adults take cover under stones and in leaf litter in the day. References: Erwin (2001), Gidaspow (1959), Data from CAS, UASM collections

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Calopachys bohnei (Korell & Frisch) 2004 BOHNE’S BEAUTIFUL THICKISH SEARCHER Calosoma bohnei Korell & Frisch, 2004:51

Distribution: Native, New World. MÉXICO – GO. Way of Life: MACROHABITAT: Uplands, 2600 – 3050 meters altitude. MICROHABITAT: Under fallen branches and fallen trees on rocky soils of volcanic origin beneath fir-oak forest (Abies guatemalensis and Quercus magnolifolia Née, Q. castanea A. DC.). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Unknown. References: Korell & Frisch (2004), Data from NMNH collection Notes: This and the following species seem to be altitudinally separated, however both species are represented (NMNH) in a series collected in July by H.H. Smith at “Omiltemi, 8000 ft.” Calopachys omiltemius (Bates) 1891 OMILTEMI BEAUTIFUL THICKISH SEARCHER Calosoma omiltemium Bates, 1891:226

Distribution: Native, New World. MÉXICO – GO. Way of Life: MACROHABITAT: Midlands to mountains, 680 – 2230 meters altitude, in montane rain and cloud forests, and subtropical mixed oak-pine forests. MICROHABITAT: On rocky soils of volcanic origin beneath forests of Quercus magnolifolia Née, Q. castanea A. DC., with some individual Pinus montezumae Lamb. that are common in oak-dominated areas. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Nocturnal, climbing trees. References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections Calopachys viridissimus (Haury) 1880 GREENISH BEAUTIFUL THICKISH SEARCHER Calosoma viridissimus Haury 1880:164

Distribution: Native, New World. MÉXICO – OA. Way of Life: MACROHABITAT: Uplands, oak forest. MICROHABITAT: Damp soil among large stones in oak forest. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: An adult was found in September. BEHAVIOR: Active in the day, running among stones and hebaceous plants at edge of road. References: Erwin (2001), Gidaspow (1959), Data from UASM collection Notes: Only two specimens are known of this species.

SPECIES ACCOUNTS / CARABIDAE

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85

Calosoma Weber 1801 CATERPILLER HUNTERS (Plates 14 and 16) Calosoma Weber, 1801:20 Callitropa Motschulsky 1865:300 [valid subgenus for native Nearctic and Neotropical species] Castrida Motschulsky, 1866:300 [valid subgenus for native Nearctic and Neotropical species] Callistriga Motschulsky, 1866:307 Calamata Motschulsky, 1866:307 Camedula Motschulsky, 1866:304 Blaptosoma Géhin, 1876:43 [valid subgenus for native Nearctic and Neotropical species] Carabosoma Géhin, 1885: xxxii [valid subgenus for native Nearctic and Neotropical species] Carabomimus Kolbe, 1895:57 [valid subgenus for native Nearctic and Neotropical species] Microcalosoma Breuning, 1927:146 Paratropa Vacher de Lapouge, 1929:3 Acampalita Vacher de Lapouge, 1929:9 Catastriga Vacher de Lapouge, 1929:9 Caludema Jeannel, 1940:68

Number of worldwide species: 84. Number of Western Hemisphere species: 55. Taxonomy: Stable. Adelphotaxon: Callisthenes Fischer von Waldheim 1820 Type species: Carabus sycophanta Linné, 1758 Distribution: Holaractic, Neotropical, Neaustral, Oriental, Australasian, Afrotropical, Native, New World (54 species), Nearctic (13 species), Introduced to the Nearctic from Europe (one species), Neotropical (41 species), shared between Nearctic and Neotropical (seven species). Several species counted as Neotropical may actually dwell at temporate altitudes. Habitat: According to Lindroth (1961), the life history of many species in North America is known (Burgess & Collins, 1917). Adults and larvae are ravenous carnivores, specializing upon catepillars and pupae of Lepidoptera, never, as far as is known, monophagously restricted to a single host. Many species are also sub-arboreal, good climbers, some even in the larval stage. Adults arrive at caterpillar outbreaks, often from long distances via flight. They are also attracted to lights. Adults of some species, however, have reduced wings and do not fly; however, these are fast runners and cover large distances in a single night in search of prey or mates. References: Erwin (2001), Burgess & Collins (1917), Gidaspow (1959, 1963), Lindroth (1961) Calosoma (Castrida) abbreviatum Chaudoir 1869 SHORT CATERPILLER HUNTER Calosoma abbreviatum Chaudoir, 1869:371

Distribution: Native, New World. BOLIVIA; COLOMBIA; ECUADOR; PERÚ. Way of Life: MACROHABITAT: Lowlands, 3 – 110 meters altitude, on hills and brush covered plains, in cultivated land (e.g. cotton fields), and in sandy areas between the coast and oakcovered foothills in Colombia. MICROHABITAT: On partly shaded sandy ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – April, and November – December. BEHAVIOR: Unknown. References: Erwin (1991, 2001), Gidaspow (1963), Data from CAS, NMNH collections

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Calosoma (Castrida) alternans (Fabricius) 1792 RECIPROCAL CATERPILLER HUNTER Carabus alternans Fabricius, 1792:146 Calosoma Laterale Dejean, 1826:199 Calosoma granulatum Perty 1830:9 Calosoma imbricatum Brullé, 1837:42 Calosoma Brulléi Blanchard, 1843:42 Calosoma armatum Reiche, 1842:377 Callistriga coxale Motschulsky, 1865:307 Calamata rugata Motschulsky, 1865:308 Calosoma Orbignyi Géhin, 1885:59

Distribution: Native, New World. ARGENTINA – BUENOS AIRES, CORRIENTES, MISIONES, “PATAGONIA”; BOLIVIA; BRAZIL; COLOMBIA; ECUADOR; GUYANA; HONDURAS; LESSER ANTILLES – ST. CROIX, ST. BARTHELEMY, DOMINICA, MARTINIQUE; MÉXICO – CS, VC; NICARAGUA; PANAMÁ; PARAQUAY; PERÚ; TRINIDAD; URUGUAY; VENEZUELA. Way of Life: MACROHABITAT: Lowlands to uplands, 28 – 1373 meters altitude. MICROHABITAT: In a variety of habitats including dense tropical dry and moist forests, deciduous scrub forest, grasslands, and cultivated fields (i.e. Sugar Cane and Cotton). DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – December. BEHAVIOR: Adults are predaceous on the Cotton Leaf Worms (Alabama argillacea (Hübner) and Spodoptera frugiperda (J.E. Smith)) in Bolivia. Adults often found flying to lights at night. References: Allen (1977), Erwin (1991, 2001), Gidaspow (1963), Data from NMNH collection Notes: In NMNH, there is a male specimen labeled Mali, W. Africa, CICP Project, August, 1987, “Nara, Acacia.” Calosoma (Carabomimus) altipeta (Jeannel) 1940 VOLCANO CATERPILLER HUNTER Carabomimus depresicollis altipeta Jeannel, 1940:229

Distribution: Native, New World. MÉXICO – MX. Way of Life: MACROHABITAT: Mountains, Volcan at Nevado de Toluca, 3200 meters altitude. MICROHABITAT: Open pine-juniper forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Jeannel (1940), Gidaspow (1959), Data from NMNH collection Calosoma (Carabosoma) angulatum Chevrolat 1834 ANGULATE CATERPILLER HUNTER Calosoma angulatum Chevrolat, 1834: no. 44 Calosoma angulicolle Chaudoir, 1869:377 Calosoma uniforme Géhin, 1885:63 Calosoma Forreri Géhin, 1885:64

SPECIES ACCOUNTS / CARABIDAE

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87

Distribution: Native, New World. COLOMBIA; COSTA RICA; GUATEMALA; HONDURAS; MÉXICO – BJ (ISLA GUADALUPE), CM, CS, GO, JA, MH, ML, MX, NA, NL, OA, PU, SL, SI, SO, TM, VC; NICARAGUA; USA – AZ, CA, NM, TX; VENEZUELA. Way of Life: MACROHABITAT: Lowlands to uplands, 60 – 1830 meters altitude, premontane moist forest, thorn scrub, oak-savannah, possibly in cultivated land. MICROHABITAT: Leaf litter in oak forest. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – November. BEHAVIOR: Often at lights at night. In Nicaragua, adults of this species prey upon the noctuid agriculture pest, Spodoptera frugiperda (J.E. Smith). References: Erwin (1991, 2001), Gidaspow (1959), Larochelle & Larivière (2003), Maes (1989), Data from NMNH, UASM collections Calosoma (Blaptosoma) anthracinum anthracinum Dejean 1831 BLACK CATERPILLER HUNTER Calosoma Anthracinum Dejean, 1831:569 Calosoma rufinum Géhin, 1885:66

Distribution: Native, New World. MÉXICO – GO. Way of Life: MACROHABITAT: Uplands, 1828 meters altitude, in montane forests of diverse and complex subtropical mixed oak-pine forests. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Calosoma (Blaptosoma) anthracinum microgonum Bates 1891 SMALL-ANGLED CATERPILLER HUNTER Calosoma laeve microgonum Bates, 1891:225

Distribution: Native, New World. MÉXICO – DF, JA, PU, VC. Way of Life: M ACROHABITAT : Uplands to mountains, 1500 – 2520 meters altitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Calosoma (Castrida) argentinensis Csiki 1927 ARGENTINE CATERPILLER HUNTER Calosoma argentinensis Csiki, 1927:11 Calosoma Antiquum Dejean, 1831:561 Calosoma belsaki Dallas, 1939:782 Calosoma cuprifulgens Dallas, 1939:784 Calosoma miniaturum Dallas, 1939:784 Calosoma ballasi Dallas, 1939:788 Calosoma pampeanum Belsak, 1939:788 Calosoma dallasi Dallas, 1939:789

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Distribution: Native, New World. ARGENTINA – BUENOS AIRES, CATAMARCA, CHACO, CÓRDOBA, CORRIENTES, ENTRE RÍOS, FORMOSA, JUJUY, LA PAMPA, LA RIOJA, MENDOZA, MISIONES, NEUQUÉN, RÍO NEGRO, SAN JUAN, SAN LUIS, SALTA, SANTA FÉ, SANTIAGO DEL ESTERO, TUCUMÁN; BOLIVIA; BRAZIL; PARAGUAY. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 770 meters altitude, in the drier regions of its range. MICROHABITAT: On the ground in xeophytic thickets and Chaco forests, dry valleys of the pampas, and in desert scrub and grassy areas on barren soil; also in pastures and cultivated land (mostly alfalfa fields). Found in great numbers on the open salt flats of central Argenitna at night. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – May, and October – December. BEHAVIOR: Adults are nocturnal, adults take cover in the day in leaf litter. Adults are predaceous on insect larvae, principally caterpillars of Colias lesbia (F.). Adults appear newly emerged in late October, early November at the latitude of Buenos Aires. Adults are attracted to lights. References: Dallas (1939), Erwin (2001), Gidaspow (1963), Data from CAS, NMNH collections Calosoma (Carabomimus) asper (Jeannel) 1940 ROUGH CATERPILLER HUNTER Carabomimus depresicollis asper Jeannel, 1940:229

Distribution: Native, New World. MÉXICO – DF, MX, ML, VC. Way of Life: MACROHABITAT: Uplands, 2438 – 3000 meters altitude, mixed coniferous forests. MICROHABITAT: Forest floor and under bark of fallen trees. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August, and November. BEHAVIOR: Adults take cover under bark and fallen trees of conifers during the day. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Calosoma (Blaptosoma) atrovirens atrovirens Chaudoir 1869 BLACKISH-GREEN CATERPILLER HUNTER Calosoma atrovirens Chaudoir, 1869:372

Distribution: Native, New World. MÉXICO – HD, SL, TM. Way of Life: MACROHABITAT: Midlands, 1135 – 1320 meters altitude. MICROHABITAT: Oak forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August, and November. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from CAS, UASM collections Calosoma (Blaptosoma) atrovirens explanaticolle Bates 1891 BROAD-NECKED CATERPILLER HUNTER Calosoma explanaticolle Bates, 1891:225 Calosoma obscurum Géhin, 1885:65

Distribution: Native, New World. MÉXICO – DF, GJ, JA, MH, OA, PU, QT, ZA.

SPECIES ACCOUNTS / CARABIDAE

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89

Way of Life: MACROHABITAT: Midlands to mountains, 1040 – 2408 meters altitude, in tropical deciduous forest, and wet pastures, MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – November. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from CAS, UASM collections Calosoma (Calosoma) aurocinctum Chaudoir 1850 GOLD-RINGED CATERPILLER HUNTER Calosoma aurocinctum Chaudoir, 1850:420 Calosoma splendidum Perbosc, 1839:261

Distribution: Native, New World. GUATEMALA; MÉXICO – CP, DU, NL, ML, OA, SL, TM, VC, YC; NICARAGUA; USA – TX. Way of Life: MACROHABITAT: Lowlands to midlands, 200 – 800 meters altitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Adults are attracted to lights at night. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH, UASM collections Notes: In NMNH, there are two specimens that may be mislabeled, one labeled Chile and the other “Tucuman” Argentina. Calosoma (Carabomimus) bolivianum Géhin 1885 BOLIVIAN CATERPILLER HUNTER Calosoma bolivianum Géhin, 1885:65

Distribution: Native, New World. BOLIVIA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1963), Data from NMNH collection Calosoma (Blaptosoma) bridgesi Chaudoir 1869 BRIDGES’ CATERPILLER HUNTER Calosoma Bridgesi Chaudoir 1869:377

Distribution: Native, New World. ARGENTINA – TUCUMÁN; BOLIVIA; CHILE. Way of Life: MACROHABITAT: Mountains, 3200 meters altitude, in Andean high desert vegetation, MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: An adult was found in October. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1963), Data from NMNH collection

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Calosoma (Carabomimus) bulleri Breuning 1943 BULLER’S CATERPILLER HUNTER Calosoma bulleri Breuning, 1943:21

Distribution: Native, New World. MÉXICO – JA. Way of Life: MACROHABITAT: Uplands, 1257 – 1600 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959) Calosoma (Blaptosoma) chihuahua Gidaspow 1959 CHIHUAHUA CATERPILLER HUNTER Calosoma chihuahua Gidaspow, 1959:284

Distribution: Native, New World. MÉXICO – CH. Way of Life: MACROHABITAT: Uplands to mountains, 2130 – 2270 meters altitude, in pine-oak woodlands, MICROHABITAT: In meadows and on lake margins. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from UASM collection Calosoma (Carabomimus) cicatricosum Chaudoir 1869 SCARRED CATERPILLER HUNTER Calosoma cicatricosum Chaudoir, 1869:374 Calosoma balli, Rotger, 1976:294

Distribution: Native, New World. MÉXICO – DF, MX. Way of Life: M ACROHABITAT : Uplands to mountains, 2166 – 2468 meters altitude. MICROHABITAT: DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June, and September – October. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH, UASM collections Calosoma (Carabomimus) costipenne Chaudoir 1869 RIBBED CATERPILLER HUNTER Calosoma costipenne Chaudoir, 1869:375

Distribution: Native, New World. MÉXICO – GO, HD, MX. Way of Life: MACROHABITAT: Mountains, 2417 – 3026 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collections

SPECIES ACCOUNTS / CARABIDAE

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91

Calosoma (Carabomimus) depressicolle Chaudoir 1869 FLAT-NECKED CATERPILLER HUNTER Calosoma depressicolle Chaudoir, 1869:375

Distribution: Native, New World. MÉXICO – JA, MH, MX. Way of Life: MACROHABITAT: Uplands to mountains, 2100 – 3200 meters altitude, in pine/ oak, pine/fir, oak/pine/madrone forests, and pastures, MICROHABITAT: On the ground in leaf litter or among grasses. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April, and July – September. BEHAVIOR: Nocturnal, adults take cover during the day under stones. References: Erwin (2001), Gidaspow (1959), Data from UASM collection Calosoma (Carabomimus) digueti (Lapouge) 1924 DIGUET’S CATERPILLER HUNTER Eutelodontum depressicolle digueti Lapouge, 1924:39

Distribution: Native, New World. MÉXICO – JA, TL, ZA. Way of Life: MACROHABITAT: Uplands to mountains, 1590 – 3048 meters altitude, oak/pine forest, scrub forest. MICROHABITAT: Found in clearings. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May, and July – August. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Jeannel (1940), Data from NMNH, UASM collections Calosoma (Carabomimus) diminutum Bates 1891 SMALL CATERPILLER HUNTER Calosoma diminutum Bates, 1891:227 Callisthenes laevissimum Casey, 1920:170

Distribution: Native, New World. MÉXICO – MX, VC. Way of Life: M ACROHABITAT : Uplands to mountains, 1733 – 2750 meters altitude. MICROHABITAT: Pastures on slopes. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal, adults take cover in the day among grasses and under debris. References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections Calosoma (Carabosoma) eremicola Fall 1910 SOLITARY CATERPILLER HUNTER Calosoma eremicola Fall, 1910:91 Calosoma rugosipennis Schaeffer, 1911:113 Calosoma hospes Casey, 1913:63 Acamegonia peregrinatrix incerta Vacher de Lapouge, 1924:38

Distribution: Native, New World. MÉXICO – BJ; USA – CA (+ SAN CLEMENTE CATALINA ISLANDS), NM, NV.

AND

SANTA

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Way of Life: MACROHABITAT: Lowlands to uplands, 100 – 1600 meters altitude, in Eastern Mojave Desert vegetation zone. MICROHABITAT: Found in high desert, as well as in fields and along roadsides. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – July, and September. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Callitropa) externum (Say) 1823 CHURCHYARD CATERPILLER HUNTER Carabus externum Say, 1823:150 Calosoma Longipenne Dejean, 1831:568

Distribution: Native, New World. CANADA – ON; USA – AL, AR, CT, DC, DE, GA, IA, IL, IN, KS, KY, LA, MA, MD, MI, MO, MS, NC, NE, NJ, NY, OH, OK, PA, SC, TN, TX, VA, VT, WI. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 1040 meters altitude, in open country. MICROHABITAT: In fallow fields and around margins of standing water. DISPERSAL ABILITIES: Macropterous, capable of flight. Occasional climber on trees. SEASONAL OCCURRENCE: Adults have been found in February, April – October, and December. BEHAVIOR: Adults are often found at lights at night and occasionally in drift material at the margin of water bodies; they are predaceous on caterpillars of the genera Melanoplus, Cirphis, Lachnosterna, Malacosoma, and Porthetria. References: Burgess & Collins (1917), Ciegler (2000), Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Carabomimus) flohri Bates 1884 FLOHR’S CATERPILLER HUNTER Carabomimus Flohri Bates, 1884:262 Calosoma lesnei Breuning, 1931:620

Distribution: Native, New World. MÉXICO – CM, JA, MH, MX. Way of Life: MACROHABITAT: Uplands to mountains, 1950 – 3200 meters altitude, in wet pine/ oak forests, pastures and grasslands, cornfields at edge of lava flow. MICROHABITAT: On the ground among grasses and in leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May and August. BEHAVIOR: Nocturnal, adults take cover in the day under wood. References: Erwin (2001), Gidaspow (1959), Jeannel (1940), Data from NMNH, UASM collections Calosoma (Calosoma) frigidum Kirby 1837 COLD-COUNTRY CATERPILLER HUNTER Calosoma frigidum Kirby, 1837:19 Calosoma frigida levettei Casey, 1897:344

Distribution: Native, New World. CANADA – AB, BC, MB, NB, NS, ON, PE, PQ, SK; USA – CO, CT, GA, IA, IL, IN, LA, MA, MD, ME, MI, MN, NC, ND, NE, NH, NJ, NV, NY, OH, PA, RI, TX, UT, VA, VT, WI, WV.

SPECIES ACCOUNTS / CARABIDAE

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Way of Life: MACROHABITAT: Lowlands to mountains, 50 – 2500 meters altitude. MICROHABITAT: On shaded moderately dry ground covered with deep leaf litter in deciduous, mixed, or coniferous forests, tree plantations (e.g., Pinus), and orchards. DISPERSAL ABILITIES: Macropterous, capable of flight. Frequent climber on trees and shrubs. SEASONAL OCCURRENCE: Adults have been found in March, and May – November; copulating pairs in May – June; gravid females in May – June; tenerals in May – September. BEHAVIOR: Diurnal and nocturnal, adults are active on trees (e.g., Abies, Picea, Populus, Quercus) and shrubs, or adults take cover under the loose bark of dead trees, in leaf litter, and under fallen trees. Adults are gregarious. Eggs are laid singly in the ground. Some females may lay eggs for two successive years during caterpillar outbreaks. Adults found overwintering after pupation in their own pupal cavity in the ground (about 5 cm deep on average) and in rotten tree-stumps. Life span can be more than 2 years. Adult are predaceous on lepidopterous caterpillars, cicadas, and locusts. Adults and perhaps larvae may play an important economic role in the control of lepidopterous caterpillars (e.g., Heterocampa, Choristoneura) outbreaks. The larva is ground-dwelling. This species is favored by the presence of large numbers of forest defoliators. Adults occasionally found at lights at night. Frequent flier, often swarming during the day. Frequent in drift material. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Castrida) fulgens Chaudoir 1869 SHINING CATERPILLER HUNTER Calosoma fulgens Chaudoir, 1869:370

Distribution: Native, New World. BRAZIL; BOLIVIA; CHILE; COLOMBIA; ECUADOR; PARAGUAY; PERÚ; URUGUAY. Way of Life: MACROHABITAT: Lowlands, sea level – 46 meters altitude, in grasslands, prairies, and savannas. MICROHABITAT: On the ground amongst herbaceous vegetation in brushy thickets. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – April, August, and October. BEHAVIOR: In Brazil, preying on “armyworm, Lep.” Adults are attracted to lights. References: Erwin (1991, 2001), Gidaspow (1963), Data from CAS, NMNH collections Calosoma (Castrida) galapageium Hope 1838 GALÁPAGOS CATERPILLER HUNTER Calosoma Galapageium Hope, 1838:130

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: SANTIAGO (JAMES). Way of Life: MACROHABITAT: Midlands, 800 – 900 meters altitude, only at the top of Isla Santiago. MICROHABITAT: Restricted to open grassy meadows. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and December. BEHAVIOR: Unknown. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from NMNH collection

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Calosoma (Carabomimus) gebieni (Breuning) 1928 GEBIEN’S CATERPILLER HUNTER Blaptosoma laeve gebieni Breuning, 1928:46

Distribution: Native, New World. MÉXICO – MH. Way of Life: MACROHABITAT: Uplands, 2198 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959) Calosoma (Carabosoma) glabratum Dejean 1831 SMOOTH CATERPILLER HUNTER Calosoma Glabratum Dejean, 1831:565 Carabosoma bolivianum Géhin, 1885:65

Distribution: Native, New World. BOLIVIA; COLOMBIA; PANAMÁ; PERÚ. Way of Life: M ACROHABITAT : Midlands to mountains, 730 – 2650 meters altitude. MICROHABITAT: In grasslands and cultivated fields. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: Unknown. References: Erwin (1991, 2001), Gidaspow (1963), Data from CAS collection Calosoma (Castrida) granatense darwinia Van Dyke 1953:10 DARWIN’S CATERPILLER HUNTER Calosoma darwinia Van Dyke, 1953:10

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: ISABELA (ALBEMARLE). Way of Life: MACROHABITAT: Lowlands, 396 meters altitude. MICROHABITAT: Desert scrub on volcanic soils. DISPERSAL ABILITIES: Brachypterous, wings slightly reduced, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in August – September. BEHAVIOR: Unknown. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from NMNH collection Calosoma (Castrida) granatense floreanum (Basilewsky) 1968 FLOREANA CATERPILLER HUNTER Castrida floreana Basilewsky, 1968:199

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: FLOREANA (CHARLES ISLAND). Way of Life: MACROHABITAT: Lowlands, sea level – 3 meters altitude. MICROHABITAT: This species is a xerophile occurring in the desert and scrub vegetation behind the dunes and beach area. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – May. BEHAVIOR: Adults occasionally found at lights at night on the beach. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from NMNH collection

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Calosoma (Castrida) granatense granatense Géhin 1885 WIDE-LINES CATERPILLER HUNTER Calosoma granatense Géhin, 1885:59 Calosoma galapagoum Howard, 1889:191 Calosoma Howardi Linell, 1898:251 Calosoma galapageium Roeschke, 1900:59

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: BALTRA (SEYMOUR), DARWIN, (CULPEPPER), EDEN, ESPANOLA (HOOD), FERNANDINA (NARBOROUGH), FLOREANA, GENOVESA (TOWER), ISABELA (ALBEMARLE), MARCHENA (BINDLOE), PINTA (ABINGDON), PINZON (DUNCAN), PITT, SAN CRISTOBAL (CHATHAM), SANTA CRUZ (INDEFATIGABLE), SANTA FÉ (BARRINGTON), SANTIGO (JAMES), SOUTH SEYMOUR. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1640 meters altitude, dry arid and lower transition vegetation belt, and lower moist forest zone. MICROHABITAT: Desender et al. (1991) wrote of this species: “One of the Calosoma species, Calosoma granatense GEHIN, 1885, occurs on virtually every island in the archipelago (cfr. Fig. 1), especially in the dry arid and lower transition vegetation belts at the lower elevations of the volcanoes. On Fernandina and on all separate volcanoes of Isabela the species also occurs at higher elevations, mostly in so-called “pampa” or meadow vegetation. Our morphometric data show that in this species the hindwings vary in size and shape (cfr. Fig. 2). As far as we know this is the first case of wing polymorphism reported for a beetle species in the Galapagos. Full-winged specimens, which are more numerous in lowland populations, mostly have functional in direct flight muscles and indeed can exhibit considerable flight activity during nights after heavy rainfall. In lowland dry arid and transition scrub vegetation, this species most probably feeds especially on lepidoptera larvae (as deduced from our few field observations of feeding beetles) and we hypothesize that the adaptive significance of light activity is mainly related to the search for new sites or trees with abundant food. An interesting case of variation in dispersal power between populations of Calosoma granatense is observed on Volcan Cerro Azul (cfr. Fig. 3), one of the highest volcanoes in the entire archipelago, reaching 1700m altitude and moreover showing an inversion in the presence of different vegetation types along its slopes (cfr. HAMANN, 1981). Because of its height, its pronounced steepness and its localization in the western part of the archipelago (with prevailing southeast winds and fog banks rolling in from the sea), this volcano shows a relatively luxuriant vegetation on nearly its complete southwestern slopes, at least during the annual rainy season. Near the coast there is a dry deciduous steppe forest, which from about 250m altitude upwards grades into a dense but low herb-meadow vegetation with mainly grasses, sedges, herbs and some ferns.The inversion zone at an elevation of about 1000 – 1100m is indicated by a somewhat dryer and more open grassland vegetation with a small shrub species becoming more common. This zone indicates the mean upper level of clouds and is succeeded by an increasingly xerophytic and shrubby vegetation higher up the volcano. From the inversion zone onwards to the top there is thus again a more patchy and shrubby vegetation, characterized as a steppe scrub. The inversion in vegetation is also indicated by several plant species, such as Opuntia cacti, which normally only occur in the dry arid zones at lower altitude of many islands, as well as the same woody species which occur at lower elevation on Cerro Azul. Calosoma granatense beetles were collected by us during 1986 along almost the entire slope of this volcano up to the rim of the caldera. Their dispersal power as indicated by wing development (cfr. Fig. 3) is high in the lower forest, much reduced in the so-called pampa-vegetation at medium altitude and higher again in the steppe scrub athigher altitute. The results of an Analysys of

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Variance (F = 15.75,p < 0.001, n=124) between the wing development in these different populations are highly significant,whereas a posteriori multiple range tests indicate the occurrence of three main groups of populations coinciding with the three major types of vegetation. We hypothesizethat wing development has been reduced at medium altitude following past emigration events by full winged ancestral individuals, because there is probably relaxed or no selection pressure acting in this open habitat to retain a high dispersal power: feeding only can take place at the soil surface level in the continuous and low vegetation (in space as well as in time); at lower altitudes as well as above the inversion zone the vegetation structure is three dimensional and complex, composed of trees and scrubs which are moreover patchily distributed; in these circumstances there is probably a continuous selection pressure promoting the retention of a higher dispersal ability. The somewhat lessened wing development of beetles from the top zone could then be the inevitable result of the higher possibility for unidirectional gene flow to occur there as compared to the lower zone, due to the higher degree of isolation. In general, this would mean that dispersal power will always become reduced in due time unless it has an adaptive significance counteracting the effects of losses due to gene flow from an area of relaxed selection.” DISPERSAL ABILITIES: Wing polymorphic, SEASONAL OCCURRENCE: Adults have been found in January – May, and September – November. BEHAVIOR: Fully-winged adults occasionally found at lights at night. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from NMNH collection Notes: There are old yet doubtful records of this species occurring in Colombia and Perú. Calosoma (Blaptosoma) haydeni haydeni G. Horn 1870 HAYDEN’S CATERPILLER HUNTER Calosoma Haydeni G. Horn, 1870:69

Distribution: Native, New World. MÉXICO – CH; USA – AZ, CO, NM, TX. Way of Life: MACROHABITAT: Mountains, 2286 meters altitude, common in fields; sometimes in beds of stone in dry creeks. MICROHABITAT: On open ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults found in April – November; females with eggs in early June and mid August; teneral adults in July. BEHAVIOR: Predatory on caterpillars and pupae at night; adults take cover in the day under debris. Adults overwinter in the ground. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from CAS, NMNH collections Calosoma (Blaptosoma) haydeni punctulicolle Bates 1891 HOLEY-NECKED CATERPILLER HUNTER Calosoma laeve punctulicolle Bates 1891:225

Distribution: Native, New World. MÉXICO – CH, CL, DF, DU, NL, SL; USA – TX. Way of Life: MACROHABITAT: Uplands to mountains, 1676 – 2350 meters altitude, in pine-oak woodlands, grasslands. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults

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have been found in May – August, and September – November. BEHAVIOR: Nocturnal, adults take cover in the day under stones. References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections Calosoma (Blaptosoma) laeve Dejean 1826 DEJEAN’S CATERPILLER HUNTER Calosoma Laeve Dejean, 1826:210 Calosoma Chevrolati Dejean 1837:25

Distribution: Native, New World. MÉXICO – CH, DF, HD, JA, NL, MX, MH, ML, PU, TM, VC. Way of Life: MACROHABITAT: Uplands, 1829 – 1950 meters altitude. MICROHABITAT: On ground near water. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – November. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections Calosoma (Carabomimus) laevigatum hoegei Breuning 1928 HÖGE’S CATERPILLER HUNTER Calosoma cicatricosum Högei Breuning, 1928:54

Distribution: Native, New World. MÉXICO – JA. Way of Life: MACROHABITAT: Uplands, 1965 meters altitude. MICROHABITAT: On ground near water. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in September. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH, UASM collections Calosoma (Carabomimus) laevigatum laevigatum Chaudoir 1869 HÖGE’S SMOOTH CATERPILLER HUNTER Calosoma laevigatum Chaudoir, 1869:374 Calosoma nitidum Géhin, 1885:66

Distribution: Native, New World. MÉXICO – DU, GJ, HD, MX, PU, VC. Way of Life: MACROHABITAT: Mountains, 2525 – 3350, meters altitude, in fir forests. MICROHABITAT: In meadows. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – June, and August. BEHAVIOR: At Salazar, MX, found “running on the plains.” References: Erwin (2001), Gidaspow (1959), Data from NMNH, UASM collections Calosoma (Castrida) leleuporum (Basilewsky) 1968 LELEUPS’ CATERPILLER HUNTER Castrida leleuporum Basilewsky, 1968:200

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: SANTA CRUZ (INDEFATIGABLE).

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Way of Life: MACROHABITAT: Midlands, 500 – 750 meters altitude, island humid fern-sedge and Sphagnum vegetation. MICROHABITAT: On moist soil beneath Miconia shrubs in humid grasslands and in Sphagnum bogs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February, April, and December. BEHAVIOR: Adults hibernate deep in the soil on damp steep slopes, up to 30 – 50 cm; they are gregarious at this time. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from CAS, NMNH collections Calosoma (Castrida) linelli Mutchler 1925 LINELL’S CATERPILLER HUNTER Calosoma Linelli Mutchler, 1925:221 Calosoma galapagoum Linell, 1889:191

Distribution: Native, New World. ECUADOR – GALÁPAGOS ISLANDS: SAN CRISTOBAL (CHATHAM). Way of Life: MACROHABITAT: Midlands, 570 meters altitude. MICROHABITAT: Mainly in open pampa habitats with some shrubs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: Unknown. References: Basilewsky (1968), Desender & de Dijn (1989), Erwin (2001), Gidaspow (1963), Data from NMNH collection Calosoma (Callitropa) macra (LeConte) 1853 LONG CATERPILLER HUNTER Callitropa macra LeConte, 1853:400

Distribution: Native, New World. MÉXICO – CL, TM; USA – AR, LA, NM, OK, TX. Way of Life: MACROHABITAT: Midlands, 1000 – 1600 meters altitude, in open country. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – November. BEHAVIOR: Adults are occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Carabosoma) marginale Casey 1897 RIMMED CATERPILLER HUNTER Calosoma marginalis Casey, 1897:340 Calosoma lugubre LeConte, 1853:400 Calosoma Lecontei Csiki, 1927:21

Distribution: Native, New World. COSTA RICA; MÉXICO – CH, ML, NL, TM; USA – AR, CA, CO, IA, IL, KS, MO, NE, NM, OK, TX. Way of Life: MACROHABITAT: Lowlands to uplands, 285 – 1830 meters altitude, on high deserts. MICROHABITAT: On partly shaded moist or wet ground with shrubby vegetation, or small trees, on prairies, pastures, and cultivated fields (e.g., cotton), and wastelands. DISPERSAL ABILITIES:

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Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – December; oviposition in July – August. BEHAVIOR: Crepuscular, adults take cover during the day. Adults found overwintering in the ground (1.2 – 15 cm deep). Life span may be up to 3 years. Adult are predaceous on caterpillars and scarabaeids. Frequent flier, sometimes swarming on warm nights to lights. References: Erwin (1991, 2001), Gidaspow (1959), Data from NMNH collection Calosoma (Carabomimus) morelianum Bates 1891 MORELOS CATERPILLER HUNTER Calosoma morelianum Bates, 1891:228

Distribution: Native, New World. MÉXICO – DF, ML, VC. Way of Life: MACROHABITAT: Mountains, 3048 meters altitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Calosoma (Carabomimus) orizabae (Jeannel) 1891 ORIZABA CATERPILLER HUNTER Carabomimus orizabae Jeannel, 1891

Distribution: Native, New World. MÉXICO – VC. Way of Life: MACROHABITAT: Uplands, 1259 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959) Calosoma (Callitropa) palmeri G. Horn 1876 PALMER’S CATERPILLER HUNTER Callitropa palmeri G. Horn, 1876:199

Distribution: Native, New World. MÉXICO – BJ (GUADALUPE ISLAND) Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1250 meters altitude, on desert island(s) off the coast of México. MICROHABITAT: Rocky pine-oak savannah (Pinus radiate D. Don and Quercus tomentella Engelm.), desert scrub with scattered palms (Erythea edulis S. Wats.) and cypress (Cupressus guadalupensis Wats.), and small tuffs of grasses, but mostly barren because of over-grazing by introduced and now feral goats. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January, and March – April. BEHAVIOR: One specimen was found at the very summit of the Island. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Note: A single specimen in CAS is labeled Isla San Benito, West Island.

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Calosoma (Carabosoma) parvicolle Fall 1910 SMALL-NECKED CATERPILLER HUNTER Calosoma parvicollis Fall, 1910:90 Calosoma prominens Bates, 1881:21 Calosoma clemens Casey, 1914:32 Calosoma pertinax Casey, 1920:163

Distribution: Native, New World. MÉXICO – BJ, SO; USA – AZ, CA, NM, NV, UT. Way of Life: MACROHABITAT: Lowlands, -86 (at Furnace Creek, Death Valley, CA) – 236 meters altitude, on deserts and sand dunes, MICROHABITAT: Open dry ground with sparse vegetation. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – July, and September. BEHAVIOR: Adults are nocturnal and diurnal, gregarious, and predaceous on caterpillars, orthopterans, and spiders. Occasionally adults found flying to lights at night. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Carabosoma) peregrinator Guérin-Méneville 1844 TRAVELING CATERPILLER HUNTER Calosoma peregrinator Guérin-Méneville, 1844:255 Calosoma carbonatum LeConte, 1862:53 Calosoma affine Bates, 1881:21 Calosoma peregrinator ingens Casey, 1913:62 Calosoma peregrinator amplipennis Casey, 1913:62 Calosoma subgracilis Casey, 1913:63 Calosoma apacheana Casey, 1913:63

Distribution: Native, New World. MÉXICO – AG, BJ, CH, DU, DF, GJ, JA, MX, MH, ML, NA, NL, PU, SI, SO, VC, ZA; USA – AZ, CA, CO, KS, NM, OK, TX, UT. Way of Life: MACROHABITAT: Lowlands to mountians, 33 – 2350 meters altitude, in thorn scrub with yucca at riparian habitats. MICROHABITAT: On open ground covered with some vegetation in grasslands, pastures, and around golf courses. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – December; copulating pairs in June; oviposition in June – July. BEHAVIOR: Adults found overwintering in cavities in the ground. Adult are predaceous on caterpillars. Occasional climber on trees, adults take cover under stones. Adults occasionally found at lights at night. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH, UASM collections Notes: One female specimen in NMNH is labeled “Bogotá, Oct. 11, 1965.”

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Calosoma (Castrida) peruviense Mandl 1971 PERÚ CATERPILLER HUNTER Calosoma peruviense Mandl, 1971:41

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Midlands, 714 meters altitude, in the Cordillera de Vilcabamba. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Unknown. References: Erwin (2001), Mandl (1971) Calosoma (Carabomimus) politum Chaudoir 1869 FURBISHED CATERPILLER HUNTER Calosoma politum Chaudoir, 1869:373

Distribution: Native, New World. MÉXICO – PU, MX, TL. Way of Life: MACROHABITAT: Mountains, 2540 – 3400 meters altitude, in pine/fir forests, sloping pastures, meadows, and pine/alder/manzanita forest edges. MICROHABITAT: Open ground, among grasses and in leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – May, July – August, and October – November. BEHAVIOR: Nocturnal, adults take cover in the day under stones. References: Erwin (2001), Gidaspow (1959), Jeannel (1940), Data from NMNH, UASM collections Calosoma (Blaptosoma) porosifrons Bates 1891 POROUS-FRONTED CATERPILLER HUNTER Calosoma porosifrons Bates, 1891:229 Calosoma laevifrons Breuning, 1931:621

Distribution: Native, New World. MÉXICO – DU. Way of Life: MACROHABITAT: Uplands to mountains, 2150 – 2800 meters altitude, in oak/pine forest. MICROHABITAT: Riparian, this species occurs at El Salto, Durango. El Salto is an old, but still active lumber mill on the 2530 meter altitude plateau and ten miles northeast is Coyotes, another mill town. The country around El Salto and Coyotes has been logged over and burned many times, but still supports a good growth of moderate sized Yellow pines (Pinus ponderosa Dougl. ex Laws.), with a ground cover of sparse grass, and in spots, dwarf Vaccinium (blueberry); several open areas become wet meadow late in the rainy season in July and are heavily covered with tall grass; some rocky hillsides support a growth of large, spreading oaks (Quercus spp.). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from CAS, NMNH, UASM collections

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Calosoma (Carabosoma) prominens LeConte 1853 PROJECTING CATERPILLER HUNTER Calosoma angulatum LeConte, 1852:199 Calosoma prominens LeConte, 1853:400

Distribution: Native, New World. MÉXICO – BJ, CH, DU, SO; USA – AZ, CA, NM, TX. Way of Life: MACROHABITAT: Lowlands to uplands, 33 – 2040 meters altititude, in acacia/ cactus scrub. MICROHABITAT: Open dry ground with sparse vegetation. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in July – October. BEHAVIOR: Adults are attracted to lights. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH, UASM collections Calosoma (Callitropa) protracta LeConte 1862 ATTRACTIVE CATERPILLER HUNTER Calosoma protractum LeConte, 1862:52 Callitropa dolens Chaudoir, 1869:376 Callitropa truncata Haury, 1885:64

Distribution: Native, New World. MÉXICO – DU, GJ, GO, JA, MH, ML, NA, PU, OA, SO; USA – AZ Way of Life: MACROHABITAT: Midlands to uplands, 1150 – 1707 meters altitude, in acacia scrub. MICROHABITAT: On the ground in high desert where cactus meets oak and mesquite thickets. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Unknown. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Calosoma (Castrida) retusum (Fabricius) 1801 BLUNT-TOOTHED CATERPILLER HUNTER Carabus retusum Fabricius, 1801:174 Calosoma laterale Kirby, 1819:379 Calosoma Bonariense Dejean, 1831:560 Calosoma patagonense Hope, 1838:129 Calosoma proseni Dallas, 1944:5 Calosoma dallasi Stévenin, 1944:13

Distribution: Native, New World. ARGENTINA – BUENOS AIRES, CATAMARCA, CÓRDOBA, ENTRE RÍOS, LA PAMPA, MENDOZA, RÍO NEGRO, SALTA, SAN LUIS, SANTA FÉ, SANTIAGO DEL ESTERO, TUCUMÁN; BOLIVIA; BRAZIL; URUGUAY. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 642 m, in forests and drier regions with sparse vegetation, and in desert scrub. MICROHABITAT: On the ground in grasslands, pastures, and cultivated areas. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in October – December, and January – March. BEHAVIOR: Unknown.

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103

References: Erwin (2001), Gidaspow (1963), Data from CAS, NMNH collections Calosoma (Castrida) rufipenne Dejean 1831 RED-WINGED CATERPILLER HUNTER Calosoma Rufipenne Dejean, 1831:567

Distribution: Native, New World. CHILE; PERÚ. Way of Life: MACROHABITAT: Lowlands to mountains, sea level – 2300 meters altitude, on the lomas, pampas, and in cultivated fields. MICROHABITAT: On the ground in grassy and desert scrub vegetation. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March and November. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1963), Data from NMNH collection Calosoma (Castrida) sayi Dejean 1826 SAY’S CATERPILLER HUNTER Calosoma Sayi Dejean, 1826:198 Calosoma armata Laporte de Castelnau, 1835:156 Calosoma alternans Jacquelin du Val, 1857:19 Calosoma sayi abdominale Géhin, 1885:58 Calosoma sayi virginica Casey, 1897:344 Calosoma alternans cuprascens Roeschke, 1900:71

Distribution: Native, New World. BELIZE; COSTA RICA; CUBA; EL SALVADOR; GUATEMALA; HISPANOLA; HONDURAS; MÉXICO – CS, NL, PU, SL, SI, SO, OA, TM, VC; NICARAGUA; PANAMÁ, PUERTO RICO; USA – AL, AR, AZ, CA, DC, FL, GA, IA, IL, IN, KS, KY, LA, MD, MN, MO, MS, NC, ND, NJ, NY, OH, OK, PA, SC, TN, TX, VA, WI. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1210 meters altitude, in tropical dry and very dry forests, cactus/acacia desert, tropical moist forests, premontane wet forests, and wet forests. MICROHABITAT: Found on the ground in open areas in cultivated lands (many various crops and in orchards), grasslands, scrub lands, along streams and rivers, and sometimes in beach drift. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE : Adults have been found in January – November. BEHAVIOR : Adults are nocturnal, taking cover in the day under wood. Larvae are mostly diurnal. Adults are predaceous on noctuid and sphingid caterpillars and pupae, chrysomelids, elaterids, silphids, and other carabids, scarabaeid larvae, some othopterans, demapterans, cicadas, pentotomids, and coreids. Adults often swarm to lights at night and are occasionally climbers on trees and shrubs. They are most active at sunset and sunrise. Adults repeatedly found at lights at night. This species is a very important beneficial insect. References: Ciegler (2000), Erwin (1991, 2001), Gidaspow (1959), Larochelle & Larivière (2003), Maes (1989), Data from NMNH, UASM collections

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Calosoma (Calosoma) scrutator (Fabricius) 1775 EXAMINOR’S CATERPILLER HUNTER Carabus scrutator Fabricius, 1775:239

Distribution: Native, New World. CANADA – ON, PQ; GUATEMALA; MÉXICO – BJ, CH, JA, MH, NA, OA, SO, VC; USA – AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, MI, MO, MN, MS, NC, NE, NH, NJ, NM, NY, OH, OK, PA, RI, SC, SD, TN, TX, VA, WI, WV; VENEZUELA. Way of Life: MACROHABITAT: Lowlands to uplands, 50 – 2110 meters altitude, in hardwood forests and dry oak/pine forests. MICROHABITAT: On the ground or climbing plants. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in January, and March – November. BEHAVIOR: Adults are predaceous on tent and noctuid caterpillars. Copulating pairs in May – August. Adults are occasional climber on plants, e.g. Helianthus tuberosa L. Adults frequently found at lights at night and in beach drift. References: Ciegler (2000), Erwin (1991, 2001), Gidaspow (1959), Data from NMNH, UASM collections Calosoma (Calosoma) splendidum Dejean 1831 SPLENDID CATERPILLER HUNTER Calsoma Splendidum Dejean, 1831:558

Distribution: Native, New World. CUBA; DOMINICAN REPUBLIC; HAITI; MÉXICO – YC; USA – FL, GA, TX. Way of Life: MACROHABITAT: Lowlands, sea level – 190 meters altitude. MICROHABITAT: In palm forests in southern Texas. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Adults occasionally found at lights at night. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Carabosoma) sponsum Casey 1897 BETROTHED CATERPILLER HUNTER Calosoma sponsa Casey, 1897:340 Calosoma parviceps Casey, 1897:341

Distribution: Native, New World. MÉXICO – BJ (ISLA SAN BENITO); USA – AZ, CA (+ SAN CLEMETE ISLAND), NM, NV, UT. Way of Life: MACROHABITAT: Lowlands to midlands, 100 – 1236 meters altitude. MICROHABITAT: Sandy clearings. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – March, May, and August. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection

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105

Calosoma (Carabosoma) striatipenne Chaudoir 1869 FURROWED-WINGED CATERPILLER HUNTER Calosoma striatipenne Chaudoir, 1869:373 Calosoma dubitatum Géhin, 1885:66

Distribution: Native, New World. MÉXICO – PU, TL, VC. Way of Life: MACROHABITAT: Mountains, 2900 meters altitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from UASM collection Calosoma (Carabomimus) striatulum Chevrolat 1835 LINED CATERPILLER HUNTER Calosoma striatulum Chevrolat, 1835: no. 165 Calosoma striolatum Géhin, 1885:66

Distribution: Native, New World. MÉXICO – GO, HD, PU, VC. Way of Life: MACROHABITAT: Mountains, 2530 – 3100 meters altitude, in pine forests, disturbed areas, and pastures. MICROHABITAT: On the ground along road edges, among grasses in fields, and in disturbed areas under debris. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May and August. Behavior: Nocturnal, adults take cover under debris. References: Erwin (2001), Gidaspow (1959), Data from NMNH, UASM collections Calosoma (Calosoma) sycophanta (Linné) 1758 AGREEABLE CATERPILLER HUNTER Carabus sycophanta Linné, 1758:414 Calosoma azureum Letzner, 1850:95 Calosoma cupreum Letzner, 1850:95 Calosoma marginatum Letzner, 1850:95 Calosoma nigrocyaneum Letzner, 1850:95 Calosoma purpureoaureum Letzner, 1850:95 Calosoma severum Chaudoir, 1850:422 Calosoma rapax Motschulsky, 1866:309 Calosoma smaragdynum G. Rossi, 1882:204 Calosoma Habelmanni Schilsky, 1888:180 Calosoma purpureipennis Reitter, 1891:257 Calosoma corvinum Heller, 1897:312 Calosoma prasinum Vacher de Lapouge, 1907:124 Calosoma solinfectum Jännichen, 1914:298 Callipara anthracinum Vacher de Lapouge, 1924:42 Calosoma Lapougei Breuning, 1927:174 Calosoma nigrosuturale Jacquet, 1930:76 Calosoma nigroaeneum Polentz, 1937:7

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Distribution: Introduced from Europe. PALEARCTIC; CANADA – NS; USA – CT, DE, MA, MD, ME, MI, NH, NJ, NM, NY, PA, RI, VT, WA. Way of Life: MACROHABITAT: Lowlands, 40 – 244 meters altitude. MICROHABITAT: On shaded moderately moist ground in deciduous forests and orchards (e.g., apple). DISPERSAL ABILITIES: Macropterous, capable of flight. Adults and larvae are frequent climbers on trees and shrubs. SEASONAL OCCURRENCE: Adults have been found in January – December but rarely after July; copulating pairs in June – July; gravid females in May. BEHAVIOR: Diurnal and nocturnal (adult and larva), adults are active on trees and shrubs (e.g. climbing Quercus trees), or on the ground. Females spend more time eating on the ground than do males. Adults and larvae are gregarious. Eggs are laid in the ground. Adults have been found overwintering in cavities in the ground (5 – 50 cm deep); some individuals remain dormant for almost 2 years. Life span is at least 2 – 4 years. Adult are predaceous on caterpillars and pupae (e.g., Lymantriidae, Thaumatopoeidae) and on coccinellids. Larvae are predaceous on caterpillars and pupae. Members of this species play an important economic role as a specific predator, in the control of the Gypsy Moth (Lymantria dispar (L.)) and Browntail Moth (Euproctis chrysorrhoea (L.)). This species is favored by Gypsy Moth outbreaks, human activities, and spring floods along large rivers. Adults frequently found at lights at night particularly in the spring. References: Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Calosoma (Castrida) trapezipenne Chaudoir 1869 DIM-WINGED CATERPILLER HUNTER Calosoma trapezipenne Chaudoir, 1869:369

Distribution: Native, New World. ARGENTINA – CATAMARCA, CÓRDOBA, LA RIOJA , MENDOZA, RÍO NEGRO, SAN JUAN, SAN LUIS. Way of Life: MACROHABITAT: Midlands to uplands, 119 – 454 meters altitude, on the western plateau of Argentina, uplands of Mendoza and San Juan Provinces. MICROHABITAT: On the ground in xerophyte forests of Córdoba and on the bare hills of Río Negro in dry brush and desert scrub. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1963), Data from CAS, IADIZA, NMNH collections Calosoma (Castrida) vagans Dejean 1831 ROAMING CATERPILLER HUNTER Calosoma Vagans Dejean, 1831:564

Distribution: Native, New World. ARGENTINA – CHUBUT, NEUQUÉN, RÍO NEGRO, “PATAGONIA”; CHILE; PERÚ. Way of Life: MACROHABITAT: Lowlands to midlands, 20 – 740 meters altitude, areas with cold winters. MICROHABITAT: On the ground in thickets and cultivated fields. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – March, July – August, and October – December. BEHAVIOR: Adults are nocturnal, taking cover in the day, e.g. in pastures under cow dung. References: Erwin (2001), Gidaspow (1963), Jeannel (1940), Data from CAS, NMNH collections

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107

Calosoma (Blaptosoma) viridisulcatum laetulum (Jeannel) 1940 PLEASANT CATERPILLER HUNTER Callitropa viridisulcatum laetulum Jeannel, 1940:216

Distribution: Native, New World. MÉXICO – JA. Way of Life: MACROHABITAT: Uplands, thorn scrub and agricultural lands, 1400 – 1650 meters altitude. MICROHABITAT: On the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – September. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from NMNH collection Calosoma (Blaptosoma) viridisulcatum viridilucens Beheim & Breuning 1943 BRIGHT GREEN CATERPILLER HUNTER Calosoma viridisulcatum viridilucens Beheim & Breuning, 1943:22

Distribution: Native, New World. MÉXICO – CM, GO, JA, NA, ZA. Way of Life: MACROHABITAT: Midlands, 1329 meters altitude, in pine forests at Atenquique, JA. MICROHABITAT: On the ground in litter of Pinus lambertiana Douglas. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: Unknown. References: Beheim & Breuning (1943), Erwin (2001), Gidaspow (1959), Data from CAS collection Calosoma (Blaptosoma) viridisulcatum viridisulcatum Chaudoir 1863 PLEASANT CATERPILLER HUNTER Calosoma viridisulcatum Chaudoir, 1863:114 Calosoma latesulcatum Oberthür, 1883: pl. 1

Distribution: Native, New World. MÉXICO – QT. Way of Life: MACROHABITAT: Midlands, 700 meters altitude. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Unknown. References: Erwin (2001), Gidaspow (1959), Data from CAS collection Calosoma (Calosoma) wilcoxi LeConte 1848 WILCOX’S CATERPILLER HUNTER Calsoma wilcoxi LeConte, 1848:346

Distribution: Native, New World. CANADA – ON, PQ; USA – AR, CA, CT, DC, DE, GA, IA, IL, IN, KS, MA, MD, MI, MN, MO, MS, NC, NE, NH, NJ, NY, OH, PA, RI, SC, TN, TX, VA, WI, WV. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 945 meters altitude. MICROHABITAT: On shaded ground in deciduous forests (e.g., Quercus, Acer), their edges, adjacent gardens and cultivated fields (e.g., pea, cotton), orchards (e.g., plum, peach) and on trees in cities. DISPERSAL ABILITIES: Macropterous, capable of flight. Frequent climber on trees and plants. SEASONAL OCCURRENCE: Adults have been found in January – December; copulating

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pairs in June; gravid females in June. BEHAVIOR: Nocturnal and diurnal, active in the sunshine on trees, or adults take cover during the day under stones or downed trees. Adults found overwintering in cavities in the ground (2.5 – 15 cm deep). Adults are able to hibernate through 2 winters and their life span is at least 2 – 3 years. Adult are predaceous on caterpillars and grasshoppers. Adults play an important economic role in the control of caterpillar (e.g., cankerworms) outbreaks. Adults occasionally found at lights at night. They are frequent flier (sometimes swarming by the hundreds to artificial lights at night). Frequent in drift material. Larvae are ground-dwelling. References: Ciegler (2000), Erwin (2001), Gidaspow (1959), Larochelle & Larivière (2003), Data from NMNH collection Carabus Linné 1758 WORM & SLUG HUNTERS (Plate 17) Carabus Linné, 1758:413 Tachypus Weber, 1801:19 [valid subgenus for introduced Palearctic species] Megodontus Solier, 1848:58 [valid subgenus for Holarctic species] Chrysocarabus Thomson, 1875:640 [valid subgenus for introduced Palearctic species] Oreocarabus Géhin, 1876: 12 [valid subgenus for native Nearctic species] Hemicarabus Géhin, 1876: 25 [valid subgenus for native Nearctic species] Archicarabus Seidlitz, 1887:6 [valid subgenus for introduced Palearctic species] Autocarabus Seidlitz, 1887:7 Tanaocarabus Reitter, 1896:135 [valid subgenus for native Nearctic species] Homoeocarabus Reitter, 1896:144 [valid subgenus for Holarctic species] Goniocarabus Reitter, 1896:150 Tmesicarabus Reitter, 1896:159 Aulonocarabus Reitter, 1896:192 [valid subgenus for Holarctic species] Diocarabus Reitter, 1896:185 [valid subgenus for native Nearctic species] Cancellocarabus Lutshnik, 1924:49 Neocarabus Hatch, 1949:144 See Lorenz (2005) for additional Old World subgnera and synonyms.

Number of worldwide species: 882. Number of Western Hemisphere species: 17. Taxonomy: Unstable, many new species yet to be described in the east Palearctic; no global synthesis. Adelphotaxon: Ceroglossus Solier 1848 Geographic status: Holarctic: Native, New World (11 species), Holarctic (one species), Introduced (six species) References: Erwin (2001), Lorenz (2005) Carabus (Tachypus) auratus Linné 1761 AURORA BOREALIS WORM & SLUG HUNTER Carabus auratus Linné, 1761:219

Distribution: Introduced from Europe. PALEARCTIC; USA – CT, MA, ME, NH, VT.

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109

Way of Life: MACROHABITAT: Lowlands, 4 – 355 meters altitude. MICROHABITAT: On warm, open preferably loamy ground in Europe, such as cultivated fields (e.g., hay, bean), meadows, pastures, gardens, parks, roadsides. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – July, and November. BEHAVIOR: Mostly diurnal, active in the sunshine. Adults found overwintering in the ground in Europe. Life span is several years, at least in captivity. Adult are predaceous on earthworms, caterpillars, insects, molluscs, and they also feed on carrion and vegetal matter in Europe. They are spring breeders. Very effective as a colonist. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961, 1985), Data from NMNH collection Notes: This species is a late introduction from Europe and has become established in the costal districts of New England and inland to Vermont. Carabus (Tachypus) cancellatus Illiger 1798 LATTICED WORM & SLUG HUNTER Carabus cancellatus Illiger, 1798:154

Distribution: Introduced, established. PALEARCTIC; USA – GA, NC, WI. Way of Life: MACROHABITAT: Lowlands, sea level – 100 meters altitude, in open country (Europe). MICROHABITAT: On open moist clayey ground, often arable, and on sandy grassey areas, as well as in forest clearings. DISPERSAL ABILITIES: North Carolina specimens in NMNH are brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal predator, but also diurnal during the reporductive period. Mating occurs mainly in May – June, teneral adults emerge in July – August. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth, et al. (1985), Data from NMNH collection Carabus (Diocarabus) chaimssonis Fischer von Waldheim 1820 FISCHER’S WORM & SLUG HUNTER Carabus chaimssonis Fischer von Waldheim, 1820: pl. 7 Carabus brachyderus Wiedemann, 1821:110 Carabus Groenlandicus Dejean, 1831:554 Carabus groenlandicus washingtoni Casey, 1920:155 Carabus rugosostrigatus Mandl, 1955:237

Distribution: Native, New World. CANADA – AB, BC, LB, MB, NT, ON, PQ, SK, YT; USA – AK, ME, NH. Way of Life: MACROHABITAT: Midlands to uplands, 600 – 2012 meters altitude. MICROHABITAT: On open or shaded moderately dry or moist, stony, rocky or gravelly ground covered with discontinuous vegetation (e.g., Empetrum), or on rich organic ground covered with leaf litter on moraines and alpine meadows and in coniferous (e.g., Pinus) and mixed forests (Populus, Picea). Above timber line in the White Mountains of NH. Also, in the vicinity of human habitations (urban and rural sites), obviously favored by human activities. DISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL

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OCCURRENCE:

Adults have been found in May – September; copulating pairs in July; teneral adults in August – September. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. BEHAVIOR: Nocturnal, adults take cover during the day under stones or among dead leaves. Adults found overwintering in their own pupal cavity. Coitus lasts at least 5 – 8 minutes. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Carabus (Tanaocarabus) finitimus Haldeman 1852 HALDEMAN’S WORM & SLUG HUNTER

Carabus finitimus Haldeman, 1852:373 Carabus Lecontei Casey, 1913:57 Carabus Caseyi Angell, 1914:75

Distribution: Native, New World. USA – KS, OK, TX. Way of Life: MACROHABITAT: Lowlands, 274 – 289 meters altitude, in bottomland forests along rivers. MICROHABITAT: Ground-dwelling on moist sandy ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March, and May – June. BEHAVIOR: Nocturnal, adults take cover during the day. Adults occasionally found at lights at night. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Carabus (Tanaocarabus) forreri forreri Bates 1882 FORRER’S WORM & SLUG HUNTER Carabus Forreri Bates, 1882:261 Carabus Townsendi Casey, 1905:160

Distribution: Native, New World. MÉXICO – DU; USA – AZ. Way of Life: MACROHABITAT: Mountains, 2286 – 2835 meters altitude, in mixed dry oak-pine forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September, and November. BEHAVIOR: Predator. Nocturnal, adults take cover during the day in under stones, fallen trees, or wood chips. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections Carabus (Tanaocarabus) forreri willi Deuve 2003 WILL’S WORM & SLUG HUNTER Carabus forreri willi Deuve, 2003:260

Distribution: Native, New World. MÉXICO – CH. Way of Life: MACROHABITAT: Uplands, 2110 meters altitude, in mixed dry oak-pine forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July.

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111

BEHAVIOR: Predator. Nocturnal, adults take cover during the day in under stones, fallen trees, or wood chips. References: Deuve (2003) Carabus (Carabus) goryi Dejean 1831 GORY’S WORM & SLUG HUNTER Carabus limbatus Say, 1825:77 Carabus Goryi Dejean, 1831:544 Carabus limbatus clarkei Blumenthal, 1958:64

Distribution: Native, New World. CANADA – ON; USA – AL, CT, DC, GA, IA, IL, IN, KY, MA, MD, MI, MN, MS, NC, NH, NJ, NY, OH, PA, RI, SC, TN, VA, VT, WI, WV. Way of Life: MACROHABITAT: Lowlands to uplands, 100 – 2030 meters altitude, in cool mosit forests. MICROHABITAT: On shaded moist ground covered with thick leaf litter in deciduous (e.g., Quercus spp., Acer spicatum Lam.) and coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir), orchards, cultivated fields (e.g., soybean) and cranberry bogs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Occasional climber on tree trunks, up to 20 m above ground. SEASONAL OCCURRENCE: Adults have been found in February – November; copulating pairs in April – May; gravid females in April; teneral adults in April, June – July, and September. BEHAVIOR: Nocturnal, adults take cover during the day among dead leaves and under stones, fallen trees, boards, and in rotten tree-stumps. Adults are gregarious. Adults found overwintering in woodlots under stones and in rotten fallen trees. Adults are predaceous on immature stages of the Gypsy Moth (Lymantria dispar (L.)) and softbodied insect larvae. References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Data from NMNH collection Carabus (Carabus) granulatus granulatus Linné 1758 GRANULATE WORM & SLUG HUNTER Carabus granulatus Linné, 1758:41 Carabus chalybaeus Voet, 1778:70 Carabus campestris M.F. Adams, 1817:297 Carabus confluens Fischer von Waldheim, 1828:187 Carabus parallelus Faldermann, 1835:19 Carabus niger Letzner, 1850:73 Carabus nigrescens Letzner, 1850:73 Carabus rufofemoratus Letzner, 1850:73 Carabus virescens Letzner, 1850:73 Carabus rubripes Géhin, 1876:15 Carabus cupreoaeneus Dalla Torre, 1877:25 Carabus nigroaeneus Dalla Torre, 1877:25 Carabus rubripes Dalla Torre, 1877:25 Carabus viridiaeneus Dalla Torre, 1877:25

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Carabus parvicollis Kraatz, 1878:129 Carabus forticostis Kraatz, 1878:130 Carabus haematomerus Kraatz, 1878:131 Carabus fulvipes Géhin, 1885:26 Carabus Wimmeli Schulz, 1900:160 Carabus expansus Vacher de Lapouge, 1905:304 Carabus crimeensis Breuning, 1933:527 Carabus granulatulus Mandl, 1955:249 Carabus atrocyaneus Lamy, 1973:71 Carabus atrocyanescens Lamy, 1973:72

Distribution: Introduced. PALEARCTIC; CANADA – BC, SK, ON, PQ; USA – MA, TX, WA. Way of Life: MACROHABITAT: Lowlands, sea level – 246 meters altitude. MICROHABITAT: Hygrophilus, on slightly shaded or open moderately dry clayish ground close or in wet meadows close to water in low deciduous forests (often stands of Alnus), swamps, along river banks and lake shores, at the borders of marshes, pools, and roadside ditches, in cultivated fields, gardens, and orchards (e.g., apple). Occasionally found in greenhouses. DISPERSAL ABILITIES : Brachypterous in North America, hence flightless thus vagility limited to walking or running. Wing-dimorphic in Europe: macropterous form capable of flight. Frequent climber on trees, tree-stumps, or low plants. SEASONAL OCCURRENCE: Adults have been found in May – July, and September – November. BEHAVIOR: Mostly nocturnal, spring breeder, usually adults take cover during the day under the loose bark of tree-stumps. Adults are gregarious in winter. Eggs are laid in cavities in the ground, larvae are found in summer months. Adults overwinter in woodlots, under the loose bark of standing tree-stumps (e.g., Alnus, Ulmus, Acer), in rotten tree-stumps, and in fallen timber. Adult are predaceous on snails, insects, and earthworms. Larvae are predaceous on snails. Markedly favored by human activities, thus effective as a colonist. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth, et al. (1985), Data from NMNH collection Carabus (Carabus) granulatus hibernicus Lindroth 1956 CELTIC WORM & SLUG HUNTER Carabus granulatus hibernicus Lindroth, 1956:7

Distribution: Introduced. PALEARCTIC; CANADA – NB, NF, NS, PM, PQ. Way of Life: MACROHABITAT: Lowlands, sea level – 200 meters altitude. MICROHABITAT: On open moist or wet, clayish ground covered with some vegetation. Along field margins, in cultivated fields (e.g., hay, potato); at borders of marshes and temporary pools, mud flats, lake shores, and in orchards. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – September, and November. BEHAVIOR: Nocturnal, adults take cover during the day under stones, pieces of wood or cardboard, and under dead fallen plants. Markedly favored by human activities, thus effective as a colonist. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1956, 1961)

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Carabus (Tanaocarabus) hendrichsi Bolívar y Pieltain 1967 HENDRICHS’ WORM & SLUG HUNTER Carabus hendrichsi Bolívar y Pieltain, 1967:156

Distribution: Native, New World. MÉXICO – NL, TM. Way of Life: MACROHABITAT: Mountains, 2300 to 3500 meters altitude, in both mixed and coniferous forests. MICROHABITAT: On the ground in forests of predominately Pinus hartwegii Lindl, P. strobiformis Engelmann, Abies vejari Marinez, and at the lower parts of the altitudinal range, Quercus spp., and at the higher part of their range, in forests of P. culmincola Anderson & Beaman. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May, July – August, and October. BEHAVIOR: Nocturnal, adults take cover during the day under fallen trees. References: Bolívar y Pieltain (1967), Erwin (2001), Data from CAS, NMNH, UASM collections Carabus (Homoeocarabus) maeander maeander Fischer von Waldheim 1820 MENDERES RIVER WORM & SLUG HUNTER Carabus maeander Fischer von Waldheim, 1820: pl. 10 Carabus incompletus Fischer von Waldheim, 1828:303 Carabus Ehrenbergi Fischer von Waldheim, 1829:368 Carabus Lapilayi Laporte de Castelnau, 1834:89 Carabus hudsonicus Motschulsky, 1866:293 Carabus maeander Simoni von Heyden, 1879:163 Carabus maeander excatenatus Kraatz, 1880:337 Carabus maeander excostatus Kraatz, 1880:338 Carabus Lecontei Géhin, 1885:27 Carabus obscuratus Géhin, 1885:26 Carabus maeander mongolicus Vacher de Lapouge, 1905:306 Carabus maeander atlanticus Vacher de Lapouge, 1925:191

Distribution: PALEARCTIC; CANADA – AB, BC, LB, MB, NB, NF, NS, NT, ON, PE, PM, PQ, SK; USA – AK, IA, IL, IN, ME, MI, MN, MT, ND, NM, NY, PA, SD, VT, WI. Way of Life: MACROHABITAT: Lowlands to midlands, 209 – 688 meters altitude. MICROHABITAT: On open somewhat moist, clayish, muddy or peaty ground covered with moderate or dense low vegetation (e.g., Carex, Typha, Sphagnum, grasses) at edges of eutrophic marshes, ponds, Carex-bogs, fens, sloughs, and pools; swamps; edges of small lakes, slow rivers, brooks, and irrigation canals; wet meadows, low open forests, and cultivated fields (e.g., carrot). DISPERSAL ABILITIES: Wing-dimorphic. Moderate runner. Occasional climber on plants. Strong burrower. Good swimmer. SEASONAL OCCURRENCE: Adults have been found in March – November; gravid females in May; teneral adults in July – August. BEHAVIOR: Diurnal and nocturnal, active in the sunshine among vegetation (especially in spring) or adults take cover on cloudy or cool days under dead plants, in and under moss, under stones, fallen trees, and pieces of cardboard. Solitary. Adults found overwintering from October and after on higher and drier ground, in the upper zone of brook borders, roadsides, and gravel pits, in vacant lots, on hills, and at the edge of woods, under well-embedded stones, in the ground at the base of trees, under the loose bark of fallen trees, and under fallen trees. Moderately favored by human activities. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

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Carabus (Archicarabus) nemoralis nemoralis Müller 1764 WOODLAND WORM & SLUG HUNTER Carabus nemoralis O.F. Müller, 1764:21 Carabus nemoralis canadensis Vacher de Lapouge, 1908:19

Distribution: Introduced. PALEARCTIC; CANADA – AB, BC, NB, NF, NS, ON, PE, PQ; USA – CA, CT, DE, ID, IL, IN, MA, ME, MI, MN, MT, NH, NJ, NV, NY, OH, OR, PA, RI, UT, VA, VT, WA, WI, WY. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1345 meters altitude. MICROHABITAT: Eurytopic, on open or shaded moderately dry, often clayish ground rich in humus covered with some vegetation in parks, gardens, lawns, pastures, cultivated fields (e.g., potato, corn, hay, bean) and along their borders, in vacant lots, backyards, docks, ski fields, gravel pits, sand pits, roadside ditches, dumps, greenhouses, basements, orchards, thickets, hedges, deciduous and mixed forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in March – December; copulating pairs in April – June; gravid females in May; teneral adults in July – September. BEHAVIOR: Mostly nocturnal, spring breeder, usually adults take cover during the day under stones, boards, stacks of lumber, dead leaves, heaps of hay, and other debris; sometimes active on cloudy days on the ground. Coitus lasts 10 – 54 minutes; eggs are laid singly in cavities in the ground. Adults found overwintering on higher and drier ground, in hedges, at the edge of woods, and in woodlots, in cavities in the ground, under stones, in leaf litter, and under the loose bark of fallen tree stumps, able to hibernate through two winters. Adult are predaceous on slugs, earthworms, caterpillars, and a small snake. Sometimes adults are found swarming on the ground under artificial lights at night. Larval are predaceous on lepidopterous caterpillars. Adults play an important role in the control of slugs, thus they are of economic importance. Markedly favored by human activities (e.g., cultivation and transportation of agricultural and horticultural produce), thus very effective as a colonist. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Carabus (Hemicarabus) serratus Say 1823 SERRATE-SHOULDER WORM & SLUG HUNTER Carabus serratus Say, 1823:77 Carabus Lineoatopunctatus Dejean, 1826:77 Carabus Tatumi Motschulsky, 1865:293 Carabus serratus vegasensis Casey, 1913:59

Distribution: Native, New World. CANADA – AB, BC, MB, NB, NF, NS, ON, PE, PQ, SK; USA – CO, CT, DC, DE, GA, IA, ID, IL, IN, KS, MA, MD, ME, MI, MN, MT, NC, ND, NE, NH, NJ, NM, OH, OR, PA, RI, SC, SD, VA, VT, WA, WI, WV, WY. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 1200 meters altitude. MICROHABITAT: Open, rather dry, gravelly or sandy ground covered with sparse vegetation. Moraines, gravel pits, sand pits, meadows, pastures, cultivated fields and their borders (e.g., wheat, fescue, barley), vacant lots, roadsides, foot of rock-falls, railway embankments, hills, forest clearings, and open forests; in sunny glades in woodlands with fireweed, Epilobium angustifolium L. DISPERSAL ABILITIES: Wing-dimorphic: Macropterous form probably capable of flight. SEASONAL OCCURRENCE: Adults

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have been found in March – November, rarely in July; copulating pairs in June; gravid females in May; teneral adults in July – September. BEHAVIOR: Mostly nocturnal, sometimes active in the spring sunshine, usually adults take cover during the day under stones, fallen trees, pieces of cardboard, heaps of hay, or dead leaves. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and gravel pits; under plant rosettes (Verbascum spp.). Markedly favored by human activities. References: Erwin (1981, 2001), Larochelle (1972, 1975), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Carabus (Tanaocarabus) sylvosus Say 1823 SILVAN WORM & SLUG HUNTER Carabus sylvosus Say, 1823:75 Carabus Lherminieri Dejean, 1826:152

Distribution: Native, New World. CANADA – ON, PQ; USA – AL, AR, CT, DC, FL, GA, IA, IL, IN, KS, MA, MD, ME, MI, MO, MN, NC, NH, NJ, NY, OK, PA, SC, TN, TX, UT, VA, WI, WV. Way of Life: MACROHABITAT: Lowlands, 50 – 1040 meters altitude. MICROHABITAT: On shaded more or less moist ground, in wooded areas such as deciduous (e.g., Quercus), mixed, and coniferous forests (e.g., Pinus), orchards (e.g., peach), and adjacent cultivated fields (e.g., soybean, potato). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – December; gravid females in autumn; teneral adults in late spring and early summer, even early July. BEHAVIOR: Nocturnal, adults take cover during the day under fallen trees, stones, and boards. Adults have been found overwintering in woodlots under fallen trees and in the ground. Adult are predaceous on lepidopterous pupae and scarabaeids. References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Carabus (Oreocarabus) taedatus agassii LeConte 1850 AGASSI’S WORM & SLUG HUNTER Carabus Agassii LeConte 1850:209 Carabus oregonensis LeConte, 1854:16 Carabus gladiator Motschulsky, 1866:285 Carabus Agassizi von Heyden, 1879:162 Carabus taedatus canadicus Roeschke, 1900:70 Carabus patulicollis Casey, 1913:57 Carabus taedatus montanicus Casey, 1913:58 Carabus stocktonensis Casey, 1920:155 Carabus taedatus coloradensis Breuning, 1933:719

Distribution: Native, New World. CANADA – AB, BC, MB, NF, NT, ON, PQ, SK, YT; USA – AK, AZ, CO, ID, MT, NM, NV, OR, UT, WY. Way of Life: M ACROHABITAT : Lowlands, mountains, subalpine and alpine zones. MICROHABITAT: On open rather dry, mostly gravelly ground covered with thin low vegetation or dead leaves on moraines, meadows, shrub-steppes, in canyons, on mountain slopes and

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cirques, in cultivated fields (e.g., wheat, barley), on sagebrush-steppes, rangelands, in open deciduous (e.g., Populus) or coniferous forests (e.g., Abies, Picea), and in burnt forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Quick to colonize recently burnt woods. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Nocturnal, adults take cover during the day under stones and fallen trees. Adults found overwintering in woodlots in rotten tree-stumps. Adult are predaceous on caterpillars. Adults may play an important economic role in the control of the Forest Tent Caterpillar (Malacosoma disstria Hubner). References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Notes: Kavanaugh (in litt.), based on a study by one of his former students, informs me that the C. taedatus complex breaks out into 4 species. Since this has not yet been published, I report here in the sense of Lindroth (1961) followed by Larochelle & Larivière (2003). Carabus (Oreocarabus) taedatus taedatus Fabricius 1787 PINE FOREST WORM & SLUG HUNTER Carabus taedatus Fabricius, 1787:196 Carabus baccivorus Fischer von Waldheim, 1820: pl. 7 Carabus seriatus Wiedemann, 1821:109 Carabus franciscanus Casey, 1913:58 Carabus bicanaliceps Casey, 1920:154 Carabus taedatus rainieri Van Dyke, 1945:108

Distribution: Native, New World. CANADA – BC; USA – AK, CA, OR, WA. Way of Life: MACROHABITAT: Lowlands, mountains, subalpine and alpine zones. MICROHABITAT: On open ground of moraines, meadows, edges of snowfields, and in the vicinity of forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: Mostly nocturnal, usually adults take cover during the day under stones, fallen trees, and dead leaves; sometimes active during the day on snow patches. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Notes: Kavanaugh (in litt.), based on a study by one of his students, informs me that the C. taedatus complex breaks out into 4 species. Since this has not yet been published, I report here in the sense of Lindroth (1961) followed by Larochelle & Larivière (2003). Carabus (Aulonocarabus) truncaticollis truncaticollis Eschscholtz 1833 BLUNT-COLLARED WORM & SLUG HUNTER Carabus truncaticollis Eschscholtz, 1833:22 Carabus lutschnikianus Basilewsky, 1937:63 Carabus truncaticollis alaskensis Basilewsky, 1937:63

Distribution: Native, New World. CANADA – NT, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands, sea level – 84 meters altitude. Meadow slopes on the tundra. MICROHABITAT: Open moderately moist, stony or muddy ground covered with meadow-

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type vegetation, dwarf birches, or willows. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August; larvae in July – August. BEHAVIOR: Larvae dwell under moss carpets. Adults are predaceous on dipterous larvae. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH collection Carabus (Megodontus) vietinghoffi vietinghoffi Adams 1812 VIETINGHOFF’S WORM & SLUG HUNTER Carabus Vietinghoffi Adams, 1812:170 Carabus Stschegolewi Poppius, 1906:17 Carabus alaskanus Obydov, 1996:85

Distribution: Native, New World. CANADA – NT, YT; USA – AK. Way of Life: MACROHABITAT: Lowlands, 12 – 500 meters altitude. MICROHABITAT: On shaded or open ground in forests (e.g., Populus, Betula, Picea) and in barren country. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October; teneral adults in August. BEHAVIOR: Noctural, adults take cover during the day under fallen trees, stones, and other debris. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Carabus (Carabus) vinctus (Weber) 1801 BOUND WORM & SLUG HUNTER Tachypus vinctus Weber, 1801:42 Carabus interruptus Say, 1823:76

Carabus ligatus Germar, 1824:6 Carabus Carinatus Dejean, 1826:80 Carabus vinctus Georgiae Csiki, 1927:185

Distribution: Native, New World. CANADA – ON; USA – AL, CT, DC, DE, FL, GA, IL, IN, MA, MD, MN, MS, NC, NJ, NY, OH, PA, RI, SC, VA, WV. Way of Life: MACROHABITAT: Lowlands, 10 – 256 meters altitude. MICROHABITAT: On shaded or half-shaded wet, clayish or muddy ground covered with dead leaves close to water in floodplain forests (e.g., Liriodendron, Liquidambar), low deciduous forests, and swamp-forests. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – December; teneral adults in July. BEHAVIOR: Nocturnal, adults take cover during the day in leaf litter, under fallen trees, fallen branches, loose bark of fallen trees, stones, and boards. Adults found overwintering in swamp-forests; in rotten tree-stumps and under the loose bark of fallen trees. Adults are able to survive winter inundation of their hibernation site. Adult are scavengers on dead fish, meat, and fowl. Adults occasionally found walking in to lights at night. References: Ciegler (2000), Erwin (1981, 2001), Larochelle & Larivière (2003), Data from NMNH collection

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Ceroglossina Lapouge 1927 NEAUSTRAL MAGNIFICENT BEETLES Ceroglossiia Lapouge, 1927:45

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable at the species level; a plethora of subspecies have been named, but not based on breeding potentials, rather color and slight morphological variation, as well as imperfectly known ranges. A thorough genomics study would greatly benefit the taxonomy of this group. Adelphotaxon: Carabina Latreille 1802. Geographic status: Neaustral. References: Erwin (2001), Jiroux (1996), Lorenz (2005) Ceroglossus Solier 1848 NEAUSTRAL MAGNIFICENT BEETLES (Plate 18) Ceroglossus Solier, 1848:59 Ceroglossa Géhin, 1876:47

Number of worldwide species: Eight. Number of Western Hemisphere species: Eight. Taxonomy: Stable at the species level; a plethora of subspecies have been named, but not based on breeding potentials. Adelphotaxon: Calosoma Weber, 1801 Type species: Carabus suturalis Fabricius, 1775 Distribution: Argentina, Chile. Habitat: Adults of this genus are found exclusively in forested or woodlot areas, even small coppice, principally in Nothofagus and Araucaria forests, woodlots, their edges, or in thickets with proximity to farms and ranches. They are dwellers of the ground, hiding themselves during the day under downed decaying trees, leaf litter, or stones. They are brachypterous and predaceous. Larvae occur in the same microhabitats as the adults. References: Erwin (2001), Jiroux (1996), Lorenz (2005) Notes: This magnificent lineage of beetles, prized by amateurs and collectors, has a variety of interesting color forms throughout it range. It is ripe for serious molecular studies that will help us understand breeding potentials and lead to a stabilization of the species taxonomy. The treatment below does not contain the most recent barrage of subspecies descriptions published in obscure journals; they will be treated in the first supplementum. Ceroglossus buqueti andestus Kraatz-Koschlau 1887 RAMPART MAGNIFICENT BEETLE Ceroglossus Buqueti andestus Kraatz-Koschlau, 1887:193 Carabus Cyaneus Faz, 1925:22 Carabus Salbachi Faz, 1925:23

Distribution: Native, New World. CHILE.

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Way of Life: MACROHABITAT: Lowlands to midlands, 500 – 1100 meters altitude, in forests and groves of Southern Beech (Nothofagus), MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March and October – November. BEHAVIOR: See description under genus, above. Adults are gregarious, up to ten individuals under a fallen tree; they take cover under dead tree trunks during the day, hunting at night on the surface. Cohabits with adults of C. chilensis seladonicus and C. magellanicus caburgansis. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti argentinensis Jiroux 1996 ARGENTINE MAGNIFICENT BEETLE Ceroglossus buqueti argentinensis Jiroux, 1996:121

Distribution: Native, New World. ARGENTINA – NEUQUÉN. Way of Life: MACROHABITAT: Midlands to uplands, 600 – 1900 meters altitude, in forests and woodlots of Nothofagus pumilio (Poepp. et Endl.) Krasser. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti arieli Jiroux 1996 ARIEL’S MAGNIFICENT BEETLE Ceroglossus buqueti arieli Jiroux, 1996:123

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 211 meters altitude, in dense forest thickets. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with adults of C. magellanicus magellanicus. References: Erwin (2001), Jiroux (1996) Ceroglossus buqueti arriagadai Jiroux 1996 ARRIAGADA’S MAGNIFICENT BEETLE Ceroglossus buqueti arriagadai Jiroux, 1996:125

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, mixed forests, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis fallaciosus. References: Erwin (2001), Jiroux (1996)

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Ceroglossus buqueti breuningi Heinz & Jiroux 2001 BURENING’S MAGNIFICENT BEETLE Ceroglossus buqueti breuningi Heinz & Jiroux, 2001:10

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 250 – 300 meters altitude, in secondary humid Araucaria forests with bamboo, also in dense stands of Nothofagus. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February, and November – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus buqueti buqueti (Laporte de Castelnau) 1835 BUQUET’S MAGNIFICENT BEETLE Carabus Buqueti Laporte de Castelnau, 1835:158 Carabus chilensis Guérin-Méneville, 1835: pl. 3 Carabus Valdiviae Hope, 1837:128 Carabus Valdiviae Gerstaecker, 1858:431 Ceroglossus Valdivianus Morawitz, 1886:439 Ceroglossus confusus Kraatz, 1887:228

Distribution: Native, New World. ARGENTINA – CHUBUT, NEUQUÉN, RÍO NEGRO; CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 859 meters altitude in thickets and coppice of Nothofagus, Gevuina, and Persea spp. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March and September – October. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis kraatzianus, C. magellanicus candens, and in the area of Mafil with C. guerini. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti chiloensis (Hope) 1838 CHILOÉ ISLAND MAGNIFICENT BEETLE Carabus chiloensis Hope, 1838:128 Carabus insularis Hope, 1838:129 Ceroglossus dorsiger Motschulsky, 1865:284 Carabus elegantissimus Reed, 1874:53 Ceroglossus Buqueti chloridus Géhin, 1885:55 Ceroglossus Buqueti Hopei Kraatz-Koschlau, 1885:420 Ceroglossus Buqueti castroensis Kraatz-Koschlau, 1886:419 Ceroglossus Monttianus Morawitz, 1886:437 Ceroglossus Buqueti punctulatus Morawitz, 1886:438 Carabus Porteri Faz, 1925:14 Carabus Buqueti Claudii Faz 1925:22 Ceroglossus buqueti penai Van Dorsselaer, 1955:88

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Distribution: Native, New World. ARGENTINA – CHUBUT, NEUQUÉN; CHILE – ISLA CHILOE. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1219 meters altitude, in forest woodlots across a broad territory, in biotopes of very different kinds, MICROHABITAT: Grounddwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti cupreus (Faz) 1925 COPPERY MAGNIFICENT BEETLE Carabus Sybarita Cupreus Faz, 1925:23 Ceroglossus psittacus Gerstaecker, 1858:425 Ceroglossus sybarita calvus Géhin, 1885: pl. 54 Ceroglossus sybarita tomentosus Kraat-Koschlau, 1885:421

Distribution: Native, New World. CHILE – ISLA CHILOE. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996, 2001) Ceroglossus buqueti deuvei Jiroux 1996 DEUVE’S MAGNIFICENT BEETLE Ceroglossus buqueti deuvei Jiroux, 1996:126

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 750 – 1300 meters altitude, in Araucaria forests [Araucaria araucana (Molina) K. Koch]. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis ficheti and C. magellanicus dolhemi. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti lepidus Kraatz-Koschlau 1891 YELLOWISH MAGNIFICENT BEETLE Ceroglossus Buqueti lepidus Kraatz-Koschlau, 1891:11 Carabus Similis Faz, 1925:22 Carabus Similis Elegans Faz, 1925:22 Carabus Similis Incertus Faz, 1925:22 Ceroglossus valdiviae inexpectatus cherquencoensis, Van Dorsselaer, 1955:90

Distribution: Native, New World. ARGENTINA – “LO. QUILLEN;” CHILE. Way of Life: MACROHABITAT: Midlands, 700 – 1350 meters altitude, in thickets of Southern Beech (Nothofagus), MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence

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flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February, and December. BEHAVIOR: See description under genus, above. Particularly on sloping hills, adults are found in biotopes with the rarer C. gloriosus. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti lorenzi Jiroux 1998 LORENZ’S MAGNIFICENT BEETLE Carabus curtus Germain, 1895:58 Ceroglossus buqueti lorenzi Jiroux, 1998:171

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1998) Ceroglossus buqueti magdalenaensis Jiroux 1996 MAGDALENA ISLAND MAGNIFICENT BEETLE Ceroglossus buqueti magdalenaensis Jiroux, 1996:129

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 10 meters altitude, coastal, extremely wet island forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus buqueti peladosus Kraatz-Koschlau 1887 PELADA MOUNTAINS MAGNIFICENT BEETLE Ceroglossus Valdiviae peladosus Kraatz-Koschlau, 1887:358 Ceroglossus Buqueti inexpectatus Kraatz-Koschlau, 1890:252 Carabus Similis Politus Faz, 1925:22

Distribution: Native, New World. ARGENTINA – CHUBUT, NEUQUÉN; CHILE. Way of Life: M ACROHABITAT : In the Cordillera Pelada, exactly where is unknown. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996)

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Ceroglossus buqueti subnitens Kraatz-Koschlau 1885 SLIGHTLY SHINY MAGNIFICENT BEETLE Ceroglossus buqueti subnitens Kraatz-Koschlau, 1885:434 Carabus Buqueti Nepotulus Faz, 1925:22

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 74 – 1000 meters altitude, in thickets of Southern Beeches (Nothofagus) and Araucaria forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus buqueti sybarita Gerstaecker 1858 LUXURIOUS MAGNIFICENT BEETLE Ceroglossus sybarita Gerstaecker, 1858:426

Distribution: Native, New World. CHILE – ISLA CHILOE. Way of Life: MACROHABITAT: Lowlands, sea level – 203 meters altitude, primary, humid, mixed forests. MICROHABITAT : Ground-dwelling at borders of lakes. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis solieri, C. suturalis ancudanus, and C. darwini darwini. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis angolicus Kraatz-Koschlau 1888 ANGOL MAGNIFICENT BEETLE Ceroglossus gloriosus chilenicus Kraatz-Koschlau, 1888:109 Ceroglossus gloriosus angolicus Kraatz-Koschlau 1888:110 Carabus Gloriosus percuprinus Faz, 1925:19 Carabus Gloriosus Subviridus Faz, 1925:19 Carabus Gloriosus Jaffueli Faz, 1925:19 Carabus Gloriosus Viridis Faz, 1925:19 Carabus Darwinié Coerulens Faz, 1925:19

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 65 – 600 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March, August, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection

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Ceroglossus chilensis chilensis (Eschscholtz) 1829 CHILEAN MAGNIFICENT BEETLE Carabus chilensis Eschscholtz, 1829:9 Ceroglossus ignitus Géhin, 1885:97 Ceroglossus Lossbergi Kraatz-Koschlau, 1887:195 Ceroglossus victoriensis Dallas, 1936:75

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 521 – 800 meters altitude, near the ocean, in thickets, south of Concepcíon, on the southern bank of Rio Itata and the northern bank of Rio BioBio, about thirty kilometres from the ocean. The topography of the region at Hualqui on the Bio Bio River varies, starting from the west with coastal lowland marshes, then crossing a low coastal mountain (dominated by Araucaria pines) range running north to south, followed by a rich fertile interior valley, and finally the Andean mountain range in the east. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March and November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis colchaguensis (Reed) 1875 COLCHAGUA MAGNIFICENT BEETLE Carabus colchaguensis Reed, 1875:225

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 800 meters altitude, in Southern Beech forests [Nothofagus alpina Poepp. & Endl. and N. dombeyi (Mirb.) Blume], MICROHABITAT: River banks in the leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus chilensis cyanicollis Kraatz 1887 BLUE-COLLARED MAGNIFICENT BEETLE Ceroglossus cyanicollis Kraatz, 1887:230 Carabus Chilensis Germanus Faz, 1925:18

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands to uplands, 600 – 2200 meters altitude, in forests and coppice of Nothofagus obliqua (Mirbel.) Bl. MICROHABITAT: Ground-dwelling on fertile, welldrained soils. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996)

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125

Ceroglossus chilensis evenoui Jiroux 1996 EVENOU’S MAGNIFICENT BEETLE Ceroglossus chilensis evenoui Jiroux, 1996:51

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands to uplands, 700 – 1800 meters altitude, in araucaria, tepa and laurel forests between the Cordillera Malla-Malla and Tolhuaca National Park, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. Cohabits with members of C. buqueti subnitens in the southern end of its range. References: Erwin (2001), Jiroux (1996) Ceroglossus chilensis fallaciosus Kraatz 1880 DECEPTIVE MAGNIFICENT BEETLE Ceroglossus chilensis fallaciosus Kraatz, 1880:56 Ceroglossus pyrilampes Morawitz, 1886:406 Ceroglossus gloriosus incertus Kraatz-Koschlau, 1888:312

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in mixed forests, small isolated thickets and coppice of Nothofagus. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. Adults are active at night and predaceous. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Notes: Found in between Vichuquen in the north and Arauco in the south. This area is heavily polluted and undergoing deforestation; conifers are replacing the Beech forests. Ceroglossus chilensis ficheti Jiroux 1996 FICHET’S MAGNIFICENT BEETLE Ceroglossus chilensis ficheti Jiroux, 1996:53

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 500 – 1300 meters altitude, in mixed forests of Nothofagus dombeyi (Mirb.) Blume and Araucaria araucana (Molina) K. Koch, At lower altitudes, Araucaria araucana drops out, but not C. c. ficheti, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with members of C. buqueti deuvei and C. magellanicus dolhemi. References: Erwin (2001), Jiroux (1996)

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Ceroglossus chilensis germaini Jiroux 1996 GERMAIN’S MAGNIFICENT BEETLE Ceroglossus chilensis germaini Jiroux, 1996:55

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 185 – 250 meters altitude, in wet and luxuriant arucaria forests between Volcano Casa Blanca and Lago Puyehue, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti chiloensis and C. darwini reedi. References: Erwin (2001), Jiroux (1996) Ceroglossus chilensis gloriosus Gerstaecker 1858 GLORIOUS MAGNIFICENT BEETLE Ceroglossus gloriosus Gerstaecker, 1858:429 Ceroglossus gloriosus carinulatus Motschulsky, 1865:284 Ceroglossus gloriosus Pradieri Géhin, 1885:55 Ceroglossus gloriosus unicostulatus Géhin, 1885:55 Ceroglossus gloriosus dorsosolutus Morawitz, 1886:403 Ceroglossus gloriosus temucensis Kraatz-Koschlau, 1890:257 Carabus Aeneovirescens Aeneus Faz, 1925:20 Carabus Aeneovirescens Faz, 1925:20 Ceroglossus chilensis gloriosus mallecoanus Vacher de Lapouge, 1928:7 Carabus diversicostatus Dallas, 1828:81 Ceroglossus chilensis Porteri Dallas, 1929:351 Ceroglossus debilicostis Dallas, 1936:74

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 4 – 933 meters altitude, in thickets, coppice and groves. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February, October, and December. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti (b. lepidus at Tolhuaca, b. buqueti at Valdivia, b. arieli at Mehuin) and with C. magellanicus (m. boeufi at Tolhuaca, m. morawitzi at Valdivia, m. magellanicus at Mehuin). References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis kraatzianus Morawitz 1886 KRAATZ’S MAGNIFICENT BEETLE Ceroglossus Kraatzianus Morawitz, 1886:432

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 43 meters altitude, in thickets and coppice of Nothofagus, Gevuina, and Persea spp., MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. C. chilensis kraatzianus

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127

adults are common and cohabit with C. buqueti buqueti and C. magellanicus candens, which also are extremely well represented. References: Erwin (2001), Jiroux (1996) Ceroglossus chilensis latemarginatus Kraatz-Koschlau 1889 TILED-MARGIN MAGNIFICENT BEETLE Ceroglossus gloriosus latemarginatus Kraatz-Koschlau, 1889:110 Ceroglossus dynastes Born, 1898:656 Ceroglossus chilensis latemarginatus splendidus Breuning, 1943:109 – 111.

Distribution: Native, New World. ARGENTINA – NEUQUÉN; CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 150 – 576 meters altitude, in coniferous forests. M ICROHABITAT : Ground-dwelling on forested slopes. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis legrandi Heinz & Jiroux 2001 LEGRAND’S MAGNIFICENT BEETLE Ceroglossus chilensis legrandi Heinz & Jiroux, 2001:2

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude, in primary forest of rather dry Nothofagus with scattered Yucca plants.. MICROHABITAT: Ground dwelling DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. References: Erwin (2001), Heinz & Jiroux (2001) Ceroglossus chilensis meridionalis Heinz & Jiroux 2001 SOUTHERN MAGNIFICENT BEETLE Ceroglossus chilensis meridionalis Heinz & Jiroux, 2001:3

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 – 500 meters altitude, in very humid Nothofagus forests. MICROHABITAT: Ground dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with C. darwini patagonensis. References: Erwin (2001), Heinz & Jiroux (2001)

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Ceroglossus chilensis mochae (Reed) 1874 MOCHA ISLAND MAGNIFICENT BEETLE Carabus Mochae Reed, 1874:54 Carabus mochae conchyliatus Géhin, 1875:56 Carabus Aeneovirescens Mochae aequicostatus Faz, 1925:20

Distribution: Native, New World. CHILE – ISLA MOCHA. Way of Life: MACROHABITAT: Lowlands, sea level – 390 meters altitude, in Araucaria araucana (Molina) K. Koch forests at the extreme southern tip of the Cordillera de Nahuelbuta - Mocha Island which rises from sea level steeply to upland forests. MICROHABITAT: Ground dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and November. BEHAVIOR: See description under genus, above. This is the only form of Ceroglossus to inhabit this isolated part of the Cordillera de Nahuelbuta. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis nigritulus Mandl 1977 MANDL’S MAGNIFICENT BEETLE Ceroglossus chilensis nigritulus Mandl, 1977:6

Distribution: Native, New World. ARGENTINA – CHUBUT. Way of Life: MACROHABITAT: Midlands, 800 meters altitude, Alerce forests, i.e. southern cypress, Fitzroya cupressoides (Molina) Johnston. MICROHABITAT: Ground dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. Cohabits with C. darwini minutissimus in the National Park, however C. c. nigritulus predominate near Lake Menendez, whereas members of C. d. minutissimus are more frequent in the remainder of the park. References: Erwin (2001), Jiroux (1996) Ceroglossus chilensis pseudopatagoniensis Heinz & Jiroux 2001 FALSE PATAGONIAN MAGNIFICENT BEETLE Ceroglossus chilensis pseudopatagoniensis Heinz & Jiroux, 2001:3

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 – 500 meters altitude, primary Nothofagus forest. MICROHABITAT: Ground dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti chiloensis and C. darwini patagoniensis. References: Erwin (2001), Heinz & Jiroux (2001)

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129

Ceroglossus chilensis seladonicus Kraatz-Koschlau 1887 MOON MAGNIFICENT BEETLE Ceroglossus gloriosus seladonicus Kraatz-Koschlau, 1887:194 Ceroglossus gloriosus hypocrita Kraatz-Koschlau, 1887:194 Carabus Aeneovirescens Ruizi Faz, 1925:20 Carabus Aeneovirescens Plagiarus Faz, 1925:20 Ceroglossus chilensis gloriosus araucanus Vacher de Lapouge, 1928:7

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 500 – 1000 meters altitude, in thickets and coppice of Nothofagus alpina Poepp. & Endl., N. pumilio (Poepp. et Endl.) Krasser, and N. dombeyi (Mirb.) Blume, etc. MICROHABITAT: In the region of Caburga, Cordillera de Lonquimaya Neltume, this subspecies is found in large groves of these southern beeches under debris on the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti andestus and C. magellanicus caburgansis. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis solieri Roeschke 1900 SOLIER’S MAGNIFICENT BEETLE Ceroglossus Solieri Roeschke, 1900:61 Ceroglossus chilensis solieri viridiobscurus Breuning, 1943:47

Distribution: Native, New World. ARGENTINA – NEUQUÉN, RÍO NEGRO; CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 203 meters altitude, in primary, wet, and impenetrable mixed forests, MICROHABITAT : Ground-dwelling. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February, October, and December. BEHAVIOR: See description under genus, above. Cohabits with members of C. buqueti chiloensis, C. suturalis puxel and C. darwini darwini; and on Isla Chiloe with C. buqueti chiloensis, C. buqueti sybarita, C. suturalis ancudanus and C. darwini darwini. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus chilensis villaricensis Kraatz-Koschlau 1885 VILLARICA MAGNIFICENT BEETLE Ceroglossus gloriosus villaricensis Kraatz-Koschlau, 1885:431 Carabus Gloriosus Villaricensis Capuccinus Faz, 1925:19 Carabus Gloriosus Darwinii Reedjuniori Faz, 1925:19

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 135 – 400 meters altitude, in a cordillera with difficult access, inside thickets and coppice. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January, June, and December. BEHAVIOR: See description

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under genus, above. Adults are rather common here often seen crossing the road. Adults take cover under debris in forested thickets. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus darwini darwini (Hope) 1837 DARWIN’S MAGNIFICENT BEETLE Carabus Darwini Hope, 1837:129 Ceroglossus indiconotus Solier, 1849:127 Ceroglossus melanopterus Gerstaecker, 1858:429 Ceroglossus Darwini bimarginatus Kraatz-Koschlau, 1886:418 Ceroglossus indiconotus disputatus Kraatz-Koschlau, 1887:228 Ceroglossus indiconotus opacus Kraatz-Koschlau, 1887:228 Ceroglossus Darwini chonchicus Kraatz-Koschlau, 1887:356 Ceroglossus darwini auroviridis Bruening, 1943:111 Ceroglossus darwini minutissimus Mandl, 1975:83

Distribution: Native, New World. CHILE – ISLA CHILOE. Way of Life: MACROHABITAT: Lowlands, sea level – 203 meters altitude, in primary, humid, mixed forests, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus darwini jirouxi Deuve 1995 JIROUX’S MAGNIFICENT BEETLE Ceroglossus darwini jirouxi Deuve, 1995:11

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 30 – 400 meters altitude, in primary forest on the Rio Manihuales. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus darwini mequignoni Heinz & Jiroux 2001 MEQUIGNON’S MAGNIFICENT BEETLE Ceroglossus darwini mequignoni Heinz & Jiroux, 2001:6

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude in primary forest. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. References: Erwin (2001), Heinz & Jiroux (2001)

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131

Ceroglossus darwini patagoniensis Breuning 1931 PATAGONIAN MAGNIFICENT BEETLE Ceroglossus darwini patagoniensis Breuning, 1931: no. 7

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 598 meters altitude. The massive primary forest in the Valle de Lago Blanco was completey burned in 1940 to raise sheep. All that is left now are grasslands and a few thickets. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus darwini reedi Morawitz 1886 REED’S MAGNIFICENT BEETLE Ceroglossus Reedi Morawitz, 1886:421

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 185 – 250 meters altitude, wet and luxuriant arucaria forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus guerini Géhin 1885 GUERIN’S MAGNIFICENT BEETLE Ceroglossus Guerini Géhin, 1885:55 Ceroglossus explanatus Breuning, 1943:24 Ceroglossus intermedius Breuning, 1943:25

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 859 meters altitude in thickets and coppice of Nothofagus, Gevuina, and Persea spp. MICROHABITAT: Ground-dwelling with densely shaded understory. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti buqueti. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus araucanus Kraatz-Koschlau 1886 AURACANA MAGNIFICENT BEETLE Ceroglossus similis araucanus Kraatz-Koschlau, 1886:148

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 74 meters altitude, in Araucaria araucana (Molina) K. Koch forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence

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flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus magellanicus boeufi Jiroux 1996 BOEUF’S MAGNIFICENT BEETLE Ceroglossus magellanicus boeufi Jiroux, 1996:87

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 500 – 1000 meters altitude, in Araucaria araucana (Molina) K. Koch forests, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis gloriosus at Victoria and Tolhuaca, and with C. buqueti lepidus at Tolhuaca. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus bouchardi Jiroux 1996 BOUCHARD’S MAGNIFICENT BEETLE Ceroglossus magellanicus bouchardi Jiroux, 1996:90

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 150 meters altitude, in mixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus caburgansis Dueve 1989 CATABURGA MAGNIFICENT BEETLE Ceroglossus darwini caburgansis Dueve, 1989:14

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 600 – 1000 meters altitude, in humid Beech forests (Nothofagus alpina Poepp. & Endl., N. pumilio (Poepp. et Endl.) Krasser, N. dombeyi (Mirb.) Blume), MICROHABITAT: Ground-dwelling near the edge of water bodies. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti andestus, and C. chilensis seladonicus. References: Erwin (2001), Jiroux (1996)

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133

Ceroglossus magellanicus candens Géhin 1885 BRIGHT MAGNIFICENT BEETLE Ceroglossus candens Géhin, 1885:96 Ceroglossus bicostulatus Kraatz-Koschlau, 1885:428 Ceroglossus Morawitzi Kraatz, 1887:236

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 43 meters altitude, in thickets and coppice of Nothofagus, Gevuina, and Persea spp., MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti buqueti and C. chilensis kraatzianus. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus davidsoni Jiroux 1998 DAVIDSON’S MAGNIFICENT BEETLE Ceroglossus magellanicus davidsoni Jiroux, 1998:170

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. Probably cohabits with C. buqueti peladosus in the forests of the Cordillera Pelada. References: Erwin (2001), Jiroux (1998) Ceroglossus magellanicus dolhemi Jiroux 1996 DOLHEM’S MAGNIFICENT BEETLE Ceroglossus magellanicus dolhemi Jiroux, 1996:91

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Midlands, 1000 – 1300 meters altitude, in mixed forests (N. dombeyi (Mirb.) Blume and Araucaria araucana (Molina) K. Koch), MICROHABITAT: Grounddwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti deuvei and C. chilensis ficheti. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus gerstaeckeri Morawitz 1886 GERSTAECKER’S MAGNIFICENT BEETLE Ceroglossus Gerstaeckeri Morawitz, 1886:429 Ceroglossus tenebriculus Kraatz-Koschlau, 1887:357 (?) Carabus Magnus Faz, 1925:11 Carabus magnus vicinus Faz, 1925:11

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Carabus magnus caputolivae Faz, 1925:11 Carabus magnus imitator Faz, 1925:11 Ceroglossus imitator Jiroux, 1996:81

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 35 – 750 meters altitude, in dense, dry, impenetrable coppices near the Cordillera de Nahuelbuta. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus magellanicus magellanicus Géhin 1885 SOUTHERN-MOST MAGNIFICENT BEETLE Ceroglossus magellanicus Géhin, 1885:56 Carabus valdiviae aerosus Faz, 1925:10 Carabus valdiviae proximus Faz, 1925:10 Carabus magnus fusco Faz, 1925:11

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 211 meters altitude, in thickets, coppice and groves. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus magellanicus newtoni Jiroux 1999 NEWTON’S MAGNIFICENT BEETLE Ceroglossus magellanicus newtoni Jiroux, 1999:211

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1999) Ceroglossus magellanicus ortrudheinzae Heinz & Jiroux 2001 ORTRUDHEINZ’S MAGNIFICENT BEETLE Ceroglossus magellanicus ortrudheinzae Heinz & Jiroux, 2001:10

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude, in primary forest with bamboo. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus

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vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and December. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis gloriosus and C. buqueti buqueti. References: Erwin (2001), Heinz & Jiroux (2001) Ceroglossus magellanicus similis Kraatz-Koschlau 1885 FAZ’S MAGNIFICENT BEETLE Ceroglossus similis Kraatz-Koschlau, 1885:435 Carabus Hyocrita Celadonicus galbinus Faz, 1925:10 Carabus Hyocrita Celadonicus aestectus Faz, 1925:10 Carabus Hyocrita Celadonicus Izquierdoi Faz, 1925:10

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus ochseni (Germain) 1895 OCHSEN’S MAGNIFICENT BEETLE Carabus Ochseni Germain, 1895:40 Ceroglossus pantherinus Deuve, 1989:57

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, 195 meters altitude, in evergreen forest with Alerce. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996) Ceroglossus speciosus Gerstaecker 1858 SHOWY MAGNIFICENT BEETLE Ceroglossus speciosus Gerstaecker, 1858:438 Carabus Speciosus Oyarzuni Faz, 1925:14

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands, sea level – 650 meters altitude, on forested slopes and hills. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996)

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Ceroglossus suturalis ancudanus Morawitz 1886 ANCUD MAGNIFICENT BEETLE Ceroglossus suturalis ancudanus Morawitz, 1886:387 Ceroglossus suturalis chiloensis Kraatz-Koschlau, 1885:435

Distribution: Native, New World. CHILE – ISLA CHILOE, MAULLIN. Way of Life: MACROHABITAT: Lowlands, sea level – 203 meters altitude, in primary, very humid, mixed dense forests, MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. Cohabits with C. chilensis solieri and C. buqueti sybarita. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus suturalis olivaceus Kraatz-Koschlau 1886 OLIVACEOUS MAGNIFICENT BEETLE Ceroglossus suturalis olivaceus Kraatz-Koschlau, 1886:420 Ceroglossus suturalis suturelevatus Kraatz-Koschlau, 1888:311 Ceroglossus suturalis speciosissimus Vacher de Lapouge, 1928:6

Distribution: Native, New World. CHILE – ISLA CHILOE. Way of Life: MACROHABITAT: Lowlands, 203 meters altitude, in primary, wet, and impenetrable mixed forest. MICROHABITAT: Ground-dwelling in thickets on escarpments. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. Cohabits with C. buqueti chiloensis. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Ceroglossus suturalis puxel Jiroux 1996 PUXEL MAGNIFICENT BEETLE Ceroglossus suturalis puxel Jiroux, 1996:139

Distribution: Native, New World. CHILE. Way of Life: MACROHABITAT: Lowlands to midlands, 300 to 900 meters altitude, in Alerce forests [Fitzroya cupressoides (Molina) Johnston]. Alerce grows on mountain slopes and lake shores. Summer average temperatures are 13 – 16°C, or higher at lower altitudes. Winter average temperatures vary from 2 – 4°C. In Chile, rainfall is as high as 6000 mm per annum. MICROHABITAT: Ground-dwelling in leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. References: Erwin (2001), Jiroux (1996), www.conifers.org

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Ceroglossus suturalis suturalis (Fabricius) 1775 SEAMED MAGNIFICENT BEETLE Carabus suturalis Fabricius, 1775:238 Carabus Reichei Guérin-Méneville, 1839:297 Carabus Dubitabilis Faz, 1925:14 Carabus Suturalis Magallanicus Faz, 1925:15 Carabus Suturalis Australis Faz, 1925:15

Distribution: Native, New World. ARGENTINA – SANTA CRUZ, TIERRA DEL FUEGO; CHILE – ISLA NAVARINO, ISLA PICTON. Way of Life: MACROHABITAT: Lowlands, sea level, in windswept thickets of Nothofagus pumilio (Poepp. et Endl.) Krasser, and other inhospitable lowland landscapes at the tip of South America. This species is the southern most occurring of the genus. MICROHABITAT: Ground-dwelling near the sea. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January, March – May, and Ocotober. BEHAVIOR: Members of this species take cover during the day under rotten fallen trees. References: Erwin (2001), Jiroux (1996), Data from NMNH collection Cychrini Laporte de Castelnau 1834 SNAIL-EATING BEETLES Cychrini Laporte de Castelnau, 1834:86

Number of worldwide genera: Six. Number of Western Hemisphere genera: Three. Taxonomy: Stable. Adelphotaxon: Carabini Latreille 1802 Geographic status: Holarctic. Behavior. Adults of this tribe stridulate, the sound louder the larger the species members. They do so by bending the head and prothorax forward, thereby scraping the inner epipleural surfaces against the lateral margin of the tergites (Roeschke, 1907). References: Erwin (2001), Lorenz (2005), (Roeschke, 1907) Cychrus Fabricius 1794 RARE SNAIL-EATING BEETLES (Plate 19) Cychrus Fabricius, 1794:440 Cychrys Fabricius, 1794:440 Kryptocychrus Cavazzuti, 1997:75 See Lorenz (2005) for additional Old World subgnera.

Number of worldwide species: 117. Number of Western Hemisphere species: 2. Taxonomy: Stable at the species level; a plethora of subspecies have been named, but not based on breeding potentials. Likely, there are many new species to be discovered in China. Adelphotaxon: Cychropsis Boileau 1901

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Type species: Carabus rostratus Fabricius, 1775 Distribution: Holarctic: Nearctic (two species), Palearctic (79 species). Habitat: According to Lindroth (1961), most species of this genus are confined to forested country. Adults are noctunral; most species hibernate as larvae or old adults, at least in part. References: Erwin (2001), Lindroth (1961), Lorenz (2005) Cychrus hemphillii hemphillii G. Horn 1878 HEMPHILL’S RARE SNAIL-EATING BEETLE Cychrus Hemphillii G. Horn, 1878:184

Distribution: Native, New World. USA – ID, UT, WY. Way of Life: MACROHABITAT: Uplands to mountains, 1676 – 3350 meters altitude, in forests or brushy forest fragments (e.g. Alnus, Philadelphicus) growing along rivers in canyons. Also, found in mixed coniferous forest with Populus and Abies. MICROHABITAT: On shaded rather dry ground with dense vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or slow running. SEASONAL OCCURRENCE: Adults have been found in May – September; teneral adults in June. BEHAVIOR: Nocturnal predator on snails and slugs. Larvae hibernate. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, NMNH collections Notes: Both forms, C. h. hemphillii and C. h. rickseckeri have representative specimens labeled Logan Pass, Utah. This area ranges from 1357 – 3050 meters altitude. Perhaps the two subspecies are altitudinally separated where their ranges overlap. Cychrus hemphillii rickseckeri LeConte 1884 RICKSECKER’S RARE SNAIL-EATING BEETLE Cychrus Rickseckeri LeConte, 1884:2

Distribution: Native, New World. CANADA – BC; USA – ID, MT, OR, UT, WA. Way of Life: MACROHABITAT: Lowlands to uplands, 500 – 1500 meters altitude, mixed and coniferous forests often along streams. MICROHABITAT: On shaded rather dry ground with some vegetation DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or slow running. SEASONAL OCCURRENCE: Adults have been found in May – August; teneral adults in June. BEHAVIOR: Nocturnal predator on snails; taking cover under stones or large fallen branches, or trunks on the ground. Larvae hibernate. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, NMNH collections Notes: See note above. Cychrus tuberculatus T.W. Harris 1839 TUBERCULATE RARE SNAIL-EATING BEETLE Cychrus tuberculatus T.W. Harris, 1839:200 Cychrus pustulosus Casey, 1905:160

Distribution: Native, New World. CANADA – BC, QC; USA – CA, OR, WA.

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Way of Life: MACROHABITAT: Lowlands to midlands, subalpine, and alpine zones, sea level – 914 meters altitude, in coniferous forests, forest clearings, or on moraine. MICROHABITAT: Shaded, rather moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – October; teneral adults in May – July. BEHAVIOR: Adults are nocturnal, taking cover in the day; they are predaceous on snails. Larvae hibernate. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus Dejean 1826 SNAIL-EATING BEETLES (Plate 20) Scaphinotus Dejean, 1826:17 Scaphinopterus Fischer von Waldheim, 1828:136 Irichroa Newman, 1838:385 Scaphonotus Agassiz, 1847:960 Analobus Gistel, 1857:90 Megalridia Casey, 1920:175 Nomaretus LeConte, 1853:399 [valid subgenus for native Nearctic species] Stenocantharis Gistel, 1857:92 [valid subgenus for native Nearctic species] Phemphus Motschulsky, 1866:312 Brennus Motschulsky, 1866:311 [valid subgenus for native Nearctic species] Pseudonomaretus Roeschke, 1907:154 [valid subgenus for native Nearctic species] Neocychrus Roeschke, 1907:197 [valid subgenus for native Nearctic species] Maronetus Casey, 1914:30 [valid subgenus for native Nearctic species] Steniridia Casey, 1924:336 [valid subgenus for native Nearctic species]

Number of worldwide species: 55. Number of Western Hemisphere species: 55. Taxonomy: Stable. Adelphotaxon: Sphaeroderus Dejean 1826 Type species: Carabus elevatus Fabricius, 1787 Distribution: Nearctic. Habitat: According to Lindroth (1961), most species of this genus are confined to forested country, many to mountainous areas. Adults are nocturnal; most species hibernate as larvae, at least in part. Adults are adapted to eating land snails, although Noonan (1967) observed adults of one species feeding on a variety of other prey items (beetles, moth larvae, berries). Notes: Numerous subspecies have been named. These need to be more fully investigated with better distributional data, as well as their population genetics. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Noonan (1967) Scaphinotus (Steniridia) aeneicollis (Beutenmüller) 1903 COPPERY-COLLARED SNAIL-EATING BEETLE Cychrus aeneicollis Beutenmüller, 1903:515 Cychrus purpuratus Beutenmüller, 1918:89

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Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Midlands to uplands, 914 – 1980 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: In southern Appalachia along streams in ravines on shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Steniridia) andrewsi amplicollis (Casey) 1920 WIDE-COLLARED SNAIL-EATING BEETLE Irichroa violacea amplicollis Casey, 1920:174 Irichroa andrewsi reflexa Casey, 1924:22

Distribution: Native, New World. USA – NC, VA. Way of Life: MACROHABITAT: Midlands to uplands, 1220 – 1524 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi andrewsi (T.W. Harris) 1839 ANDREWS’ SNAIL-EATING BEETLE Cychrus Andrewsii T.W. Harris, 1839:195

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Lowlands, 140 – 156 meters altitude, in deciduous and mixed forests (Quercus, Pinus, Cornus, and Carya spp.). MICROHABITAT: On shaded moist ground along the banks of woodland streams on the southern Appalachia piedmont. A specimen collected in Virginia in 1943 is labeled “Oak-Chestnut.” DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August, October and December. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark, in rotten wood, or in tree hollows; adults are gregarious in autumn. Adults eat berries and snails. Regular climber on tree trunks while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi barksdalei (Valentine) 1936 BARKSDALE’S SNAIL-EATING BEETLE Steniridia andrewsi barksdalei Valentine, 1936:230

Distribution: Native, New World. USA – NC.

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Way of Life: MACROHABITAT: Midlands to uplands, 914 – 1676 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi darlingtoni (Valentine) 1935 DARLINGTON’S SNAIL-EATING BEETLE Steniridia andrewsi darlingtoni Valentine, 1935:356

Distribution: Native, New World. USA – NC, TN. Way of Life: MACROHABITAT: Midlands to uplands, 610 – 1768 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moist ground on ridges and in ravines. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – October. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi germari (Chaudoir) 1861 GERMAR’S SNAIL-EATING BEETLE Cychrus Germari Chaudoir, 1861:495

Distribution: Native, New World. USA – KY, TN, VA. Way of Life: MACROHABITAT: Lowlands to midlands, 450 – 1220 meters altitude in the Cumberland Mountains, in mixed forests (e.g., Quercus spp., Acer spicatum Lam., Osytra spp.) on slopes and in ravines with a dense growth of eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbiense Boursault and R. maximum L.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – July, and September – October. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark of fallen trees. Adults are predators on snails. Regular climber on fallen tree trunks and cave walls while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH, UASM, collections Scaphinotus (Steniridia) andrewsi montanus (Valentine) 1935 MOUNTAIN SNAIL-EATING BEETLE Steniridia andrewsi montana Valentine, 1935:350

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Midlands to uplands, 1060 – 1530 meters altitude, in mixed forests (e.g., Quercus spp., Acer spicatum Lam., Osytra spp.) on slopes and in ravines with a dense growth of

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eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbiense Boursault and R. maximum L.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On shaded moist somewhat rocky ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – June, and August – September. BEHAVIOR: Nocturnal, taking cover in the day under loose tree bark of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi mutabilis (Casey) 1920 CHANGELING SNAIL-EATING BEETLE Irichroa mutabilis Casey, 1920:173 Irichroa mutabilis longicollis Casey, 1920:173 Irichroa mutabilis modulatus Casey, 1920:174

Distribution: Native, New World. USA – KY, OH, PA, WV. Way of Life: MACROHABITAT: Lowlands, 370 – 500 meters altitude, in mixed forests (e.g., Quercus spp., Acer spicatum Lam.). MICROHABITAT: On shaded moist ground covered with leaf litter, stones, and fallen trees. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – December. BEHAVIOR: Nocturnal, taking cover in the day in rotten fallen trees; adults are gregarious in winter. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi nantahalae (Valentine) 1936 NANTAHALA SNAIL-EATING BEETLE Steniridia andrewsi nantahalae Valentine, 1936:228

Distribution: Native, New World. USA – NC, SC. Way of Life: MACROHABITAT: Midlands to uplands, 670 – 1829 meters altitude, in forests. MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Nocturnal, taking cover in the day in rotten tree stumps and fallen trees, occasionally under moss carpets. Frequent climber on tree stumps and fallen trees while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNH, UASM collections Scaphinotus (Steniridia) andrewsi parvitarsalis (Valentine) 1935 SMALL-FOOTED SNAIL-EATING BEETLE Steniridia andrewsi parvitarsalis Valentine, 1935:354

Distribution: Native, New World. USA – GA, NC, TN. Way of Life: MACROHABITAT: Midlands, 610 – 1220 meters altitude, in forests. MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless

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143

thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – October. BEHAVIOR: Nocturnal, taking cover in the day under loose bark of tree stumps and fallen trees. Frequent climber on tree stumps and fallen trees while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi saludae (Valentine) 1936 HEALTHY SNAIL-EATING BEETLE Steniridia andrewsi saludae Valentine, 1936:229

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Midlands, 1142 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1936), Data from NMNH collection Scaphinotus (Steniridia) andrewsi waldensius (Valentine) 1935 WALDEN RIDGE SNAIL-EATING BEETLE Steniridia andrewsi waldensia Valentine, 1935:357

Distribution: Native, New World. USA – TN. Way of Life: MACROHABITAT: Lowlands to midlands, 450 – 914 meters altitude, in mixed forest. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935, 1936), Data from NMNH collection Scaphinotus (Neocychrus) angulatus (T.W. Harris) 1839 ANGULATE SNAIL-EATING BEETLE Cychrus angulatus T.W. Harris, 1839:200 Scaphinotus angulatus maritimus Van Dyke, 1924:5

Distribution: Native, New World. CANADA – BC; USA – OR, WA. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1067 meters altitude, in temperate coniferous rainforest (Pseudosuga). MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Occasionally climbing on tree trunks. SEASONAL OCCURRENCE: Adults have been found in April – December. BEHAVIOR: Nocturnal, taking cover in the day under old fallen trees and in leaf litter, sometimes in rotten tree trunks. Adults are predaceous on snails. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, UASM collections

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Scaphinotus (Stenocantharis) angusticollis (Fischer von Waldheim) 1823 NARROW-COLLARED SNAIL-EATING BEETLE Cychrus angusticollis Fischer von Waldheim, 1823: pl. 46 Pemphus angusticollis nigripennis Roeschke, 1907:167 Scaphinotus angusticollis olympiae Van Dyke, 1944:5

Distribution: Native, New World. CANADA – BC, QC; USA – AK, CA, OR, WA. Way of Life: MACROHABITAT: Lowlands to midlands, subalpine zone, sea level – 1387 meters altitude, in forests (coniferous, mixed, and deciduous), and cultivated fields MICROHABITAT: Shaded moist ground often along streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – November. Teneral adults in May – July, and September. BEHAVIOR: Nocturnal, gregarious (up to 50 individuals in or under a single log). Adult are predaceous on snails, slugs, earthworms, and spiders. Taking cover during the day mostly in rotten fallen trees and treestumps, under the loose bark and moss of dead standing and fallen trees, in hollow tree roots, also under fallen trees and boards, among dead leaves and other debris. Frequent climber on shrubs, tree-stumps, and fallen trees up to 20 m above ground while foraging. Adults have been found over wintering in old tree-stumps and fallen trees. Sometimes found in and around human settlements. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections Scaphinotus (Neocychrus) behrensi (Roeschke) 1907 BEHRENS’ SNAIL-EATING BEETLE Neocychrus Behrensi Roeschke, 1907:199 Scaphinotus behrensi malkini Van Dyke, 1944:13

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1170 meters altitude, in temperate coniferous rainforest (Pseudosuga). MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Occasionally climbing on tree trunks. SEASONAL OCCURRENCE: Adults have been found in May – July, and September. BEHAVIOR: Nocturnal, taking cover in the day under old fallen trees and in leaf litter, sometimes in rotten tree trunks. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, UASM collections Scaphinotus (Nomaretus) bilobus (Say) 1823 BILOBED SNAIL-EATING BEETLE Cychrus bilobus Say, 1823:73

Distribution: Native, New World. CANADA – MB, NB, NS, ON, PQ; USA – IL, KS, MA, ME, MI, MN, MO, NE, NH, NY, OH, VT, WI. Way of Life: MACROHABITAT: Lowlands, subalpine zone, 220 – 430 meters altitude, in coniferous (e.g., Picea, Abies) and mixed forests (e.g., Alnus, Acer, Betula), and their edges, also in clearings and

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roadsides. MICROHABITAT: Shaded or open moist or wet ground covered with deep leaf litter or moss, often along rivers, lakes, and the seashore, and in gravel pits surrounded by spruce forests as well as in Sphagnum-bogs and fens. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. Teneral adults in July and September. BEHAVIOR: Nocturnal, taking cover during the day under dead leaves, fallen trees, stones, or the loose bark of fallen trees. Frequent climber and good burrower while foraging. Larval habitat: in leaf litter. Adult are predaceous on snails. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Brennus) bullatus Van Dyke 1924 INFLATED SNAIL-EATING BEETLE Scaphinotus subtilis bullatus Van Dyke, 1924:3 Scaphinotus subtilis grandis Van Dyke, 1924:4

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands to mountains, 304 – 2286 meters altitude, in coniferous and mixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March, June – July, and September – November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Van Dyke, (1924), Data from CAS, NMNH collections Scaphinotus (Nomaretus) cavicollis (LeConte) 1859 CONCAVE-COLLARED SNAIL-EATING BEETLE Nomaretus cavicollis LeConte, 1859:3

Distribution: Native, New World. USA – AR, KS, MO, OK. Way of Life: MACROHABITAT: Lowlands, 164 – 353 meters altitude, in coniferous forests (Pinus). MICROHABITAT: On shaded moderately moist ground on tree-covered stony knolls. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – June, and August – December. BEHAVIOR: Nocturnal, taking cover during the day under bark of fallen trees. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections Scaphinotus (Brennus) cordatus (LeConte) 1853 HEART-SHAPED THORAX SNAIL-EATING BEETLE Cychrus cordatus LeConte, 1853:399 Brennus cordatus vernicatus Casey, 1920:183 Brennus cordatus rufitaris Casey, 1920:184

Distribution: Native, New World. USA – CA.

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Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 800 meters altitude, in coastal mixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March, May – September, and November – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH collections Scaphinotus (Brennus) crenatus (Motschulsky) 1859 NOTCHED SNAIL-EATING BEETLE Cychrus crenatus Motschulsky, 1859:161 Cychrus striatus LeConte, 1859:69 Brennus gentilis Casey, 1897:322 Brennus productus Casey, 1914:29 Brennus montereyensis Casey, 1920:177

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowland to midlands, sea level – 1295 meters altitude, in forests. MICROHABITAT: Along stream banks. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – October, and December. BEHAVIOR: Nocturnal, taking cover in the day under stones and fallen trees. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Scaphinotus (Brennus) cristatus (T.W. Harris) 1839 RIDGED SNAIL-EATING BEETLE Cychrus cristatus T.W. Harris, 1839:200 Cychrus reticulates Motschulsky, 1850:90 Brennus basalis Casey, 1897:311 Brennus duplicatus Casey, 1897:312

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1150 meters altitude, in mixed and coniferous forests (e.g. Pseudosuga). MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – December, teneral adults in May – June. BEHAVIOR: Larvae hibernate. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections

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Scaphinotus (Nomaretus) debilis alpinus (Beutenmüller) 1903 ALPINE SNAIL-EATING BEETLE Nomaretus debilis alpinus Beutenmüller, 1903:512

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Uplands, 1500 – 2030 meters altitude, in forests on Mt. Mitchell. MICROHABITAT: On shaded moderately moist ground rich in leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Occasionally climbing on tree trunks. SEASONAL OCCURRENCE: Adults have been found in May – June, and August – September. BEHAVIOR: Nocturnal, taking cover in the day under stones, loose bark of trees, and in leaf litter. References: Erwin (2001), Darlington (1931), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Nomaretus) debilis debilis (LeConte) 1853 WEAKLY SNAIL-EATING BEETLE Nomaretus debilis LeConte, 1853:399

Distribution: Native, New World. USA – GA, NC, SC. Way of Life: MACROHABITAT: Uplands, 1580 – 1740 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moderately moist ground rich in leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: Nocturnal, taking cover in the day in deep leaf litter under stones, fallen trees, and loose bark of trees. Occasional climber while foraging. Larvae found in deep leaf litter. Larvae hibernate. References: Erwin (2001), Darlington (1931), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Scaphinotus) elevatus coloradensis Van Dyke 1907 COLORADO SNAIL-EATING BEETLE Scaphinotus elevatus coloradensis Van Dyke, 1907:141

Distribution: Native, New World. CANADA – MB; USA – CO, IA, MN, NE, NM, SD. Way of Life: MACROHABITAT: Lowlands and mountain forests (Picea, Populus) along streambeds; also on the prairies, in cultivated fields (e.g. corn). MICROHABITAT: On shaded or open moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February, and April – August. BEHAVIOR: Nocturnal, taking cover in the day under stones, fallen trees, and other debris. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1961), Van Dyke (1938), Data from NMNH collection Notes: See below under the nominate subspecies.

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Scaphinotus (Scaphinotus) elevatus elevatus (Fabricius) 1787 ELEVATED THORAX SNAIL-EATING BEETLE Cychrus elevatus Fabricius, 1787:198

Distribution: Native, New World. USA – AL, CT, DE, FL, LA, MA, MD, ME, MS, NC, NH, NJ, NY, PA, RI, SC. Way of Life: MACROHABITAT: Lowland forests. MICROHABITAT: Shaded or open areas, on moderately moist ground, often in cultivated fields, meadows, and clay pits. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – December, teneral adults in April – May. BEHAVIOR: Nocturnal. According to Larochelle & Larivière (2003), “taking cover during the day under stones, fallen tree-stumps, fallen trees, the loose bark of trees and tree-stumps, dead leaves, boards, in animal burrows, rotten fallen trees and fallen trees, and scarabaeid (Xylorictes) borings at the base of trees. Gregarious in winter (up to 30 individuals in the same shelter).” References: Erwin (2001), Erwin (field notes), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Notes: In the very large NMNH collection of Scaphinotus (Scaphinotus) elevatus (Fabricius), intergrades between the named subspecies (with the exception of S. e. neomexicanus Van Dyke) are so numerous that I seriously doubt that real subspecies exist. This is a large project for genomics. I have retained the traditional subspecies here and used the data from Larochelle & Larivière (2003) plus that from the NMNH collections where possible. Scaphinotus (Scaphinotus) elevatus flammeus Haldeman 1844 FIERY SNAIL-EATING BEETLE Scaphinotus flammeus Haldeman, 1844:54 Cychrus dilatatus LeConte, 1853:398

Distribution: Native, New World. USA – AR, IA, IL, IN, KS, MO, NE, OH, TN. Way of Life: MACROHABITAT: Lowlands in deciduous forests, often along rivers. MICROHABITAT: On shaded moist ground. TLE found many adults under driftwood at the margin of a small pond in Kansas, in open range country. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – November, teneral adults in May – June. Larvae hibernate. BEHAVIOR: Nocturnal, taking cover in the day in leaf litter and under driftwood. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Notes: See above under the nominate subspecies. Scaphinotus (Scaphinotus) elevatus lengi Van Dyke 1938 LENG’S SNAIL-EATING BEETLE Scaphinotus elevatus lengi Van Dyke, 1938:122

Distribution: Native, New World. USA – VA. Way of Life: MACROHABITAT: Lowlands, known only from the Dismal Swamp in southwestern Virginia. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless

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thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Nocturnal, taking cover in the day in leaf litter and under logs and branches. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Notes: See above under the nominate subspecies. Scaphinotus (Scaphinotus) elevatus neomexicanus Van Dyke 1924 NEW MEXICO SNAIL-EATING BEETLE Scaphinotus elevatus neomexicanus Van Dyke, 1924:1

Distribution: Native, New World. USA – NM. Way of Life: MACROHABITAT: Mountains, 2440 meters altitude, in coniferous forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke 1907 PERFORATE SNAIL-EATING BEETLE Scaphinotus elevatus tenebricosus Roeschke, 1907:141

Distribution: Native, New World. USA – NC, SC, VA. Way of Life: MACROHABITAT: Lowlands and mountains in open forests. MICROHABITAT: On shaded wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – June, and October, teneral adults in May. BEHAVIOR: Nocturnal, taking cover in the day under loose bark of fallen trees. Occasional climber while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Notes: See above under the nominate subspecies. Scaphinotus (Nomaretus) fissicollis (LeConte) 1853 CLEFT-COLLARED SNAIL-EATING BEETLE Nomaretus fissicollis LeConte, 1853:399

Distribution: Native, New World. USA – AR, IA, IL, KS, MO, MN. Way of Life: MACROHABITAT: Lowlands, 83 – 386 meters altitude, in deciduous (Quercus, Carya) and coniferous (Pinus) forests. MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – June, and August – November, teneral adults in April and May. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections

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Scaphinotus (Steniridia) guyotii (LeConte) 1866 GUYOT’S SNAIL-EATING BEETLE Cychrus Guyotii LeConte, 1866:363 Cychrus Angelli Beutenmüller, 1918:89 Scaphinotus confusus Darlington, 1931:146

Distribution: Native, New World. USA – NC, TN, VA, WV. Way of Life: MACROHABITAT: Midlands to uplands, 1060 – 2030 meters altitude, at higher altitudes in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir), at lower altitudes in mixed and deciduous forests (e.g., Quercus spp., Acer spicatum Lam.), on slopes and in ravines with a dense growth of eastern hemlock (Tsuga canadensis (L.) Carr.), rhododendron (R. catawbiense Boursault and R. maximum L.) and mountain laurel (Kalmia latifolia L.). MICROHABITAT: On cool, shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – September. BEHAVIOR: Nocturnal, taking cover in the day under loose bark or wood chips. Adults are predaceous on snails. Occasional climber while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Stenocantharis) hatchi Beer 1971 HATCH’S SNAIL-EATING BEETLE Scaphinotus hatchi Beer, 1971:257

Distribution: Native, New World. USA – OR. Way of Life: MACROHABITAT: Uplands, 1650 – 1707 meters altitude, in coniferous forests, (Tsuga mertensiana (Bong.) Sarg.). MICROHABITAT: On shaded moist ground with sparse vegetation (e.g. Vaccinium scoparium Leiberg), most abundant in broad depressions of the region with more moisture. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, taking cover in the day under old fallen trees, loose bark of fallen trees, and stones. Occasional climber while foraging. References: Beer (1971), Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Scaphinotus (Maronetus) hubbardi (Schwarz) 1895 HUBBARD’S SNAIL-EATING BEETLE Nomaretus Hubbardi Schwarz, 1895:272

Distribution: Native, New World. USA – NC, TN, VA. Way of Life: MACROHABITAT: Uplands, 1524 – 1829 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). MICROHABITAT: On shaded moist ground with deep leaf litter and branches. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March, and May – October. BEHAVIOR: Nocturnal, taking cover in the day in leaf litter, under branches, and under loose bark of fallen trees. Occasional climber while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Data from CAS, NMNH collections

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Scaphinotus (Maronetus) imperfectus (G. Horn) 1860 IMPERFECT SNAIL-EATING BEETLE Nomaretus imperfectus G. Horn, 1860:569

Distribution: Native, New World. USA – MD, NC, OH, PA, VA, WV. Way of Life: MACROHABITAT: Midlands, 1036 meters altitude, in deciduous and coniferous forests, and in thickets (e.g. Rhododendron) along streams. MICROHABITAT: On shaded moist ground with rich leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – November. BEHAVIOR: Nocturnal, taking cover in the day in leaf litter, under stones, and under loose bark of fallen trees. Occasional climber while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Maronetus) incompletus (Schwarz) 1895 INCOMPLETE SNAIL-EATING BEETLE Nomaretus incompletus Schwarz, 1895:271

Distribution: Native, New World. USA – KY, VA. Way of Life: MACROHABITAT: Midlands, ca. 500 meters altitude, in mixed evergreen forest. MICROHABITAT: Unknown. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Unknown. References: Erwin (2001), Data from NMNH collection Scaphinotus (Nomaretus) infletus Allen & Carlton 1988 FORGOTTEN SNAIL-EATING BEETLE Scaphinotus infletus Allen & Carlton, 1988:132

Distribution: Native, New World. USA – AR. Way of Life: MACROHABITAT: Midlands, 570 – 627 meters altitude. MICROHABITAT: Grounddwelling in moist, well shaded areas. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – July; teneral adults in late May. BEHAVIOR: Nocturnal, taking cover in the day under mediumsized flat rocks on a steep slope along a secondary (unpaved) road about 5m from the road. References: Carleton (pers. comm.), Erwin (2001), Larochelle & Larivière (2003), Will (pers. comm.) Notes: Chris Carleton (pers. comm.) provided the following: “All were from a limited area in Newton County, Ark. in the core of the Arkansas Ozarks, vicinity of Buffalo National River. The area is topographically diverse, ranging from perhaps 300 – 700 m elevations, with dramatic bluffs and steep hillsides. Geology is a mix of limestone/ dolomite with prominent sandstone cliffs and escarpments. Forest covers are predominantly mature deciduous, with mixed shortleaf pine on high points. Land use: pasture/chicken farms/protected forests in the National River and Ozark National Forests, which is where the specimen localities are located. Lost Valley is rich beech maple woodland in a valley with a cave at one end. Alum Cove and the area south of Mt. Judea are both within the Ozark National Forest and likewise are in areas where nice, cool, moist forests dominate. The former has a hiking trail through such an area to a natural bridge that I suspect is where that paratype came from because that is where everybody goes. Apparently

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Scaphinotus infletus is sympatric with Scaphinotus cavicollis and S. fissicollis, though I don’t see any indication in our collection that any of them were collected at the same localities.” Scaphinotus (Brennus) interruptus (Ménétriés) 1843 MÉNÉTRIÉS’ SNAIL-EATING BEETLE Cychrus interruptus Ménétriés, 1843:54 Cychrus constrictus LeConte, 1853:398 Cychrus dissolutus Schaum, 1863:72 Brennus politus Casey, 1897:330 Brennus sinuatus Casey, 1897:330 Brennus corpulentus Casey, 1897:331 Brennus integer Casey, 1914:29 Brennus parvulicollis Casey, 1920:176 Brennus Beringi Casey, 1920:179 Brennus procerus Casey, 1920:179

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1220 meters altitude, in forests. MICROHABITAT: On sterile sandy ground at edge of rivers and streams, as well as in deciduous forests (e.g., Quercus), coniferous forests (e.g., Pinus, Pseudotsuga), open grassy knolls, gardens, and in cultivated fields. Shaded or open areas on moderately moist ground usually covered with some vegetation. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – July, and September – December, teneral adults in June and December. BEHAVIOR: Nocturnal, taking cover during the day under the loose bark of trees, clods of earth, fallen branches, boards, stones, and in fallen trees. Adults are gregarious; they are predaceous on snails, slugs, and sow bugs. References: Erwin (2001), Gidaspow (1968), Kavanaugh, (1977), Larochelle & Larivière (2003), Weber & Kavanaugh. (1992), Data from NMNH, UASM collections Scaphinotus (Brennus) johnsoni Van Dyke 1924 JOHNSON’S SNAIL-EATING BEETLE Scaphinotus johnsoni Van Dyke, 1924:3 Scaphinotus klahowyae Perrault, 1973:47

Distribution: Native, New World. CANADA – BC; USA – WA. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 975 meters altitude, forests (Tsuga) and subalpine zone in the Olympic Temperate Rainforest. MICROHABITAT: On shaded moist ground with dense vegetation (e.g. mosses, Oxalis, Polystictum). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – May, and July – August. BEHAVIOR: Nocturnal, taking cover in the day under and in rotten fallen trees. Adults are gregarious. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Lindroth (1961), Data from NMNH, UASM collections

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Scaphinotus (Scaphinotus) kelloggi (Dury) 1912 KELLOGG’S SNAIL-EATING BEETLE Cychrus Kelloggi Dury, 1912:104

Distribution: Native, New World. USA – NM. Way of Life: MACROHABITAT: Uplands, 1861 meters altitude, in canyons with streams or rivers. MICROHABITAT: Moist ground on riverbanks. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July – August. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Nomaretus) liebecki Van Dyke 1936 LIEBECK’S SNAIL-EATING BEETLE Scaphinotus liebecki Van Dyke, 1936:40

Distribution: Native, New World. USA – AR, LA, TX. Way of Life: MACROHABITAT: Lowlands, 73 – 108 meters altitude, in pine/oak forest. MICROHABITAT: One specimen was near a temporary marsh. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April, June, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections Scaphinotus (Steniridia) lodingi lodingi (Valentine) 1935 LÖDING’S SNAIL-EATING BEETLE Steniridia lodingi Valentine, 1935:364

Distribution: Native, New World. USA – AL. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 914 meters altitude, in forests, and occasionally in caves. MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April, and June – November. BEHAVIOR: Occasional climber on cave walls while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH, UASM collections Scaphinotus (Steniridia) lodingi obscurus (Valentine) 1935 OBSCURE SNAIL-EATING BEETLE Steniridia lodingi obscura Valentine, 1935:366

Distribution: Native, New World. USA – AL. Way of Life: MACROHABITAT: Lowlands, 250 meters altitude, forests. MICROHABITAT: On shaded wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking

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or running. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: Nocturnal, taking cover in the day under wood and in leaf litter. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Neocychrus) longiceps Van Dyke 1924 LONG-HEADED SNAIL-EATING BEETLE Scaphinotus longiceps Van Dyke, 1924:5

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands, 182 meters altitude, in coastal temperate coniferous rainforest (Sequoia, Pseudosuga) and mixed forests. MICROHABITAT: On shaded moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May, July, August, and October – December. BEHAVIOR: Nocturnal, taking cover in the day under fallen trees, in leaf litter and in rotten tree trunks. Occasional climber on tree trunks while foraging. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from CAS, NMNH collections Notes: This species is known from midlands in northern CA, however, only one locality given on labels is precise enough to determine altitude (that lowland one which is given above). Scaphinotus (Scaphinotus) macrogonus Bates 1891 BATES’ SNAIL-EATING BEETLE Scaphinotus macrogonus Bates, 1891:229 Scaphinotus horni Van Dyke, 1938:116

Distribution: Native, New World. MÉXICO – DU. Way of Life: MACROHABITAT: Midlands to mountains, 1320 – 2590 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in September. BEHAVIOR: See description under genus, above. References: Erwin (2001), Van Dyke (1938), Data from CAS, NMNH collections Scaphinotus (Pseudonomaretus) mannii Wickham 1919 MANN’S SNAIL-EATING BEETLE Scaphinotus Mannii Wickham 1919:170

Distribution: Native, New World. USA – OR, WA. Way of Life: MACROHABITAT: Lowlands, 268 – 438 meters altitude, in forests fragments and scrub in canyons and along rivers; at Wawawai, WA, with a large stand of Holodiscus discolor (Pursh) Maxim. MICROHABITAT: Shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – July, October – November, copulating pairs in June, and teneral adults in March – May. BEHAVIOR: Nocturnal, taking cover during the day in cavities under

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stones, often on canyon slopes. Larval habitat similar to adult in burrows dug in the ground. Adults found over wintering in ground crevices. Gregarious in small groups under large stones. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Brennus) marginatus (Fischer von Waldheim) 1820 MARGINED SNAIL-EATING BEETLE Cychrus marginatus Fischer von Waldheim, 1820: pl. 7 Brennus marginatus Fulleri G. Horn, 1878:179 Cychrus marginatus gracilis Géhin, 1885:76 Brennus cupripennis Casey, 1897:334 Brennus insularis Casey, 1897:334 Brennus confusus Casey, 1897:336 Brennus marginatus fallax Roeschke, 1907:174 Brennus columbianus Casey, 1920:180 Brennus marginatus montanicus Casey, 1920:182 Brennus marginatus wrangelli Casey, 1920:182 Brennus oregonus Casey, 1920:182

Distribution: Native, New World. CANADA – AB, BC, QC; USA – AK, CA, ID, MT, OR, WA, WY. Way of Life: MACROHABITAT: Lowlands to uplands, subalpine and alpine zones, sea level – 2042 meters altitude, in coniferous (e.g., Pseudotsuga, Pinus), mixed, and deciduous forests, MICROHABITAT: Shaded or open areas on moist ground. According to Larochelle & Larivière (2003) often along streams, lakes, the seashore, and canyons, in forest clearings, woodlots, and cultivated fields, spruce/fir forest, Alnus thickets, in aspen scrub vegetation, snow-patches (above timber line), and in supratidal meadows and open coastal tundra (Aleutian Islands). Members of this species are also found in vicinity of human habitations (urban and rural sites, e.g., gardens). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in January – December, teneral adults in March to early June, and late July in AK. BEHAVIOR: Nocturnal, taking cover during the day in rotten fallen trees and tree-stumps, under stones, under the loose bark of fallen trees, in hollow tree roots, in leaf litter and moss; gregarious (up to 50 individuals in the same shelter). Excellent climber (on fallen trees and tree-stumps) while foraging. Larval habitat: under stones. Adults found over wintering in fallen trees, half rotten fallen trees and tree-stumps, and in crevices in the ground. Adult are predaceous on snails and slugs. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from NMNH, UASM collections Notes: This species, as presently understood, is markedly variable across its extensive geographic and altitudinal range. It is ripe for an intense genomics study. Scaphinotus (Pseudonomaretus) merkelii (G. Horn) 1890 MERKEL’S SNAIL-EATING BEETLE Sphaeroderus Merkelii G. Horn, 1890:71 Cychrus idahoensis Webb, 1901:133

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Distribution: Native, New World. CANADA – BC; USA – ID, MT. Way of Life: MACROHABITAT: Lowlands to uplands, 700 – 1675 meters altitude, in mixed forests. MICROHABITAT: On shaded rather dry ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – August, teneral adults in late June and late July. BEHAVIOR: Nocturnal, taking cover in the day under bark (e.g. fallen Abies trunk). References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections Scaphinotus (Scaphinotus) mexicanus Bates 1882 MEXICAN SNAIL-EATING BEETLE Scaphinotus mexicanus Bates, 1882:320

Distribution: Native, New World. MÉXICO – DU. Way of Life: MACROHABITAT: Uplands to mountains, 1790 – 2835 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June and August. BEHAVIOR: Adults occasionally found walking to lights at night. References: Erwin (2001), Van Dyke (1938), Data from CAS, NMNH collections Scaphinotus (Brennus) obliquus (LeConte) 1868 OBLIQUE SNAIL-EATING BEETLE Cychrus obliquus LeConte, 1868:61 Brennus convergens Casey 1897:326 Brennus opacicollis Casey 1897:327 Brennus sculptipennis Casey 1897:327

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Midlands to uplands, 637 – 1768 meters altitude, in forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Brennus) oreophilus (Rivers) 1890 MOUNTAIN LOVING SNAIL-EATING BEETLE Cychrus oreophilus Rivers, 1890:111 Scaphinotus oreophilus Hoppingi Roeschke, 1907:183 Brennus oreophilus humeralis Casey, 1914:30

Distribution: Native, New World. USA – CA, NV, UT. Way of Life: MACROHABITAT: Lowlands to midlands, 10 – 1388 meters altitude. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to

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walking or running. SEASONAL OCCURRENCE: Adults have been found in January, and March – November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH collections Scaphinotus (Scaphinotus) parisianus Allen & Carlton 1988 OZARK SNAIL-EATING BEETLE Scaphinotus parisianus Allen & Carlton, 1988:130

Distribution: Native, New World. USA – AR. Way of Life: MACROHABITAT: Midlands, 840 – 915 meters altitude in the Ouachita Mountains. MICROHABITAT: Ground-dwelling in moist deciduous forest leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – June, and October. BEHAVIOR: Nocturnal. See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Will (pers. comm.) Scaphinotus (Scaphinotus) petersi biedermani Roeschke 1907 BIEDERMAN’S SNAIL-EATING BEETLE Scaphinotus Biedermani Roeschke, 1907:571

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands to mountains, 1765 – 2560 meters altitude, in coniferous forests. MICROHABITAT: On shaded moist ground on slopes. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, taking cover in the day under stones and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Scaphinotus) petersi catalinae Van Dyke 1924 CATALINA MOUNTAINS SNAIL-EATING BEETLE Scaphinotus catalinae Van Dyke, 1924:2

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Mountains, 2315 – 2745 meters altitude, in coniferous forests. MICROHABITAT: On shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Nocturnal, taking cover in the day under stones and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH, UASM collections

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Scaphinotus (Scaphinotus) petersi corvus (Fall) 1910 RAVEN SNAIL-EATING BEETLE Cychrus corvus Fall, 1910:89

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands to mountains, 1980 – 2745 meters altitude, coniferous forests. MICROHABITAT: On shaded moist ground along brooks. DISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, taking cover in the day under stones and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH, UASM collections Scaphinotus (Scaphinotus) petersi grahami Van Dyke 1938 GRAHAM’S SNAIL-EATING BEETLE Scaphinotus grahami Van Dyke, 1938:107

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Midlands to uplands, 828 – 2135 meters altitude, in coniferous and mixed forests. M ICROHABITAT : On shaded moist ground. D ISPERSAL ABILITIES : Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September. BEHAVIOR: Nocturnal, taking cover in the day under stones and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Scaphinotus) petersi kathleenae Ball 1966 KAY’S SNAIL-EATING BEETLE Scaphinotus petersi kathleenae Ball, 1966:714

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Mountains, 2405 – 2622 meters altitude, in mixed forests, predominately Douglas Fir (Pseudotsuga taxifolia (Poir) Britton), yellow pine (Pinus ponderosa Douglas), and Gambel’s Oak (Quercus gambelii Nuttall). MICROHABITAT: On shaded moist, but well-drained, loamy ground underlain by coarse gravel, particularly on steep slopes. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in August. BEHAVIOR: Nocturnal, taking cover in the day under fallen trees, bits of fallen trees, and in leaf litter. References: Ball, (1966), Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections

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Scaphinotus (Scaphinotus) petersi petersi Roeschke 1907 PETERS’ SNAIL-EATING BEETLE Scaphinotus Petersi Roeschke, 1907:137

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands to mountains, 1524 – 2347 meters altitude, coniferous forest. MICROHABITAT: On shaded moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: Nocturnal, taking cover in the day under fallen trees and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Brennus) punctatus (LeConte) 1859 PUNCTATE SNAIL-EATING BEETLE Cychrus punctatus LeConte, 1859:69 Cychrus mimus G. Horn, 1874:20 Brennus gravidus Casey, 1897:317 Brennus catenulatus Casey, 1897:324

Distribution: Native, New World. MÉXICO – BJ; USA – CA. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1920 meters altitude, in forests. MICROHABITAT: On shaded or open moderately moist or dry ground along lake shores forests, and streams, and in dry fields. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – October, and December, teneral adults in early June. BEHAVIOR: Nocturnal, taking cover in the day under stones, fallen trees, and dead leaves. Gregarious. Adults eat snails, live and dead caterpillars, berries, and other beetles. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Noonan (1967), Data from NMNH collection Scaphinotus (Pseudonomaretus) regularis (LeConte) 1884 MEASURED SNAIL-EATING BEETLE Cychrus regularis LeConte, 1884:2

Distribution: Native, New World. CANADA – BC; USA – ID, WA. Way of Life: MACROHABITAT: Midlands, 700 – 1000 meters altitude, in forests. MICROHABITAT: Shaded, moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – August, and October – November, teneral adults in late spring to early summer. BEHAVIOR: Nocturnal, taking cover in the day under stones, fallen trees, and dead leaves. Larval habitat: under fallen trees. Adults found over wintering in ground crevices. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH collection

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Scaphinotus (Pseudonomaretus) relictus (G. Horn) 1881 RILICTUAL SNAIL-EATING BEETLE Cychrus relictus G. Horn, 1881:188

Distribution: Native, New World. CANADA – AB, BC; USA – CA, ID, MT, WA. Way of Life: MACROHABITAT: Midlands, 648 – 1111 meters altitude, forests (deciduous, mixed, and coniferous), occasionally in human areas such as gardens, and on yellow starthistle rangeland. MICROHABITAT: Shaded, moderately moist ground with deep humus layer. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – November; copulating pairs in August – September; teneral adults in May – July, mostly in late June. BEHAVIOR: Nocturnal, taking cover during the day under stones and fallen trees. Larval habitat: under stones. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections Scaphinotus (Steniridia) ridingsii intermedius Valentine 1935 RIDINGS’ INTERMEDIATE SNAIL-EATING BEETLE Scaphinotus ridingsi intermedia Valentine, 1935:368

Distribution: Native, New World. USA – VA. Way of Life: MACROHABITAT: Midlands, 775 meters altitude, in forests. MICROHABITAT: Shaded, moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: Nocturnal, taking cover during the day. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935) Scaphinotus (Steniridia) ridingsii monongahelae Leng 1917 MONONGAHELA SNAIL–EATING BEETLE Scaphinotus monongahelae Leng, 1917:36 Irichroa tenuiceps Casey, 1920:172

Distribution: Native, New World. USA – PA, WV. Way of Life: MACROHABITAT: Lowlands, 290 – 372 meters altitude, in coniferous forests (e.g., Picea), often in gorges. MICROHABITAT: Shaded, moderately moist, stony ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in May – October, teneral adults in mid-July. BEHAVIOR: Nocturnal, taking cover during the day under fallen trees. Occasional climber (on tree trunks) while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Steniridia) ridingsii ridingsii (Bland) 1863 RIDINGS’ SNAIL-EATING BEETLE Cychrus Ridingsii Bland, 1863:353

Distribution: Native, New World. USA – VA.

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Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 915 meters altitude, forests in riparian habitats. MICROHABITAT: Found in rocky, moist sites in the shade along streams and in and around steep rocky outcroppings near the river. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in June – October, teneral adults in early July. BEHAVIOR: Nocturnal, taking cover during the day in deep leaf litter and under stones. Adult are predaceous on snails; they have been observed mating in September. Frequent climber on stones and boulders while foraging. References: Erwin (1981, 2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Brennus) riversi (Roeschke) 1907 RIVERS’ SNAIL-EATING BEETLE Brennus oreophilus Riversi Roeschke, 1907:183

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Midlands to mountains, 914 – 2804 meters altitude, forests. MICROHABITAT: Shaded, moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Nocturnal, taking cover during the day under stones and fallen trees. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH collections Scaphinotus (Brennus) rugiceps incipiens (Casey) 1897 CASEY’S SNAIL-EATING BEETLE Brennus incipiens Casey, 1897:313

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands to uplands, 24 – 1550 meters altitude, in mixed and coniferous forest MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – September. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH collections Scaphinotus (Brennus) rugiceps rugiceps (G. Horn) 1872 WRINKLE-HEADED SNAIL-EATING BEETLE Cychrus rugiceps G. Horn, 1872:143 Brennus porcatus Casey, 1897:328 Brennus compositus Casey, 1897:332 Brennus rugiceps congener Casey, 1914:28

Distribution: Native, New World. USA – CA, OR.

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Way of Life: MACROHABITAT: Lowlands to uplands, 75 – 1880 meters altitude, mixed and coniferous forests (e.g., Pinus). MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – August, and October – December. BEHAVIOR: Nocturnal, taking cover during the day under stones and fallen trees. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections Scaphinotus (Maronetus) schwarzi (Beutenmüller) 1913 SCHWARZ’S SNAIL-EATING BEETLE Nomaretus Schwarzi Beutenmüller, 1913:139

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Uplands, 1765 – 1980 meters altitude, in forests. MICROHABITAT: Shaded areas with moderately moist ground rich in leaf litter, humus, and moss, and on grassy balds. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. Strong burrower. SEASONAL OCCURRENCE: Adults have been found in June, and August – September. BEHAVIOR: Nocturnal, taking cover during the day in leaf litter, humus, and under large stones well embedded in moss. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Scaphinotus) snowi parkeri Van Dyke 1938 ROESCHKE’S SNAIL-EATING BEETLE Scaphinotus parkeri Van Dyke, 1938:101

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands to mountains, 2134 – 2591 meters altitude, in coniferous forests. MICROHABITAT: Shaded areas on moist ground, along brooks and near springs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: Nocturnal, taking cover during the day under stones and fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Scaphinotus) snowi roeschkei Van Dyke 1907 ROESCHKE’S SNAIL-EATING BEETLE Scaphinotus Roeschkei Van Dyke, 1907:135

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Mountains, 2896 meters altitude. MICROHABITAT: Grounddwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection

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Scaphinotus (Scaphinotus) snowi snowi (LeConte) 1881 SNOW’S SNAIL-EATING BEETLE Cychrus Snowi LeConte, 1881:71 Scaphinotus snowi parkeri Van Dyke, 1938:101

Distribution: Native, New World. USA – AZ, CO, NM, UT. Way of Life: MACROHABITAT: Uplands to mountains, 1675 – 2745 meters altitude, in coniferous forests. MICROHABITAT: Shaded areas on moist ground, along brooks and near springs. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: Nocturnal, taking cover during the day under stones and fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Brennus) striatopunctatus (Chaudoir) 1844 PORE-WINGED SNAIL-EATING BEETLE Cychrus striatopunctatus Chaudoir, 1844:476 Brennus decipiens Casey, 1897:316 Brennus subdepressus Casey, 1920:177

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands to uplands, sea level – 1524 meters altitude, mostly in deciduous forests (e.g., Quercus), but also in canyons and on grassy knolls. MICROHABITAT: Shaded or open areas on moderately moist, sandy ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in January – July, and October – December. BEHAVIOR: Nocturnal, taking cover during the day in rotten fallen trees, under fallen trees, or under the loose bark of fallen trees. Frequent climber on fallen trees while foraging. Adults are gregarious. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, UASM collections Scaphinotus (Brennus) subtilis (Schaum) 1863 SLENDER SNAIL-EATING BEETLE Cychrus subtilis Schaum, 1863:72

Distribution: Native, New World. USA – CA. Way of Life: MACROHABITAT: Lowlands to uplands, 89 – 1981 meters altitude, near bodies of water. MICROHABITAT: Adults are found on wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – August, and October – December. BEHAVIOR: Nocturnal, taking cover during the day. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from CAS, NMNH, UASM collections

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Scaphinotus (Maronetus) tenuis (Casey) 1914 THIN SNAIL-EATING BEETLE Maronetus tenuis Casey, 1914:31

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Uplands, above 1500 meters altitude, in forests. MICROHABITAT: Shaded areas on moderately moist ground rich in leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – July, and September. BEHAVIOR: Nocturnal, taking cover during the day in leaf litter and humus. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Steniridia) tricarinatus (Casey) 1914 THREE-RIDGED SNAIL-EATING BEETLE Irichroa aeneicollis tricarinata Casey, 1863:353

Distribution: Native, New World. USA – NC, SC, TN. Way of Life: MACROHABITAT: Midlands to uplands, 914 – 1829 meters altitude, in forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir, Betula alleghaniensis Britton, Betula alleghaniensis Britt.). MICROHABITAT: Shaded areas with moist ground, often in ravines along streams. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – October. BEHAVIOR: Nocturnal, taking cover during the day under moss carpets. Occasional climber on the base of tree trunks while foraging. At least some adults hibernate (full adult from April collection). References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH, UASM collections Scaphinotus (Scaphinotus) unicolor (Fabricius) 1787 ONE-COLOR SNAIL-EATING BEETLE Cychrus unicolor Fabricius, 1787:198 Scaphinotus heros T.W. Harris, 1839:196, new synonymy Scaphinotus grandis Gistel, 1857:52, n. dubium Scaphinotus Hunteri Crotch, 1873:5 Scaphinotus Shoemakeri Leng, 1914:143, new synonymy Scaphinotus elevatus floridanus Leng, 1915:564, new synonymy

Distribution: Native, New World. USA – AL, AR, DC, FL, GA, IN, KY, MD, MO, MS, OH, PA, SC, TN, VA. Way of Life: MACROHABITAT: Lowlands to midlands, 4 – 840 meters altitude, in forests, often along streams and near springs. MICROHABITAT: Found in and around rocky outcroppings in the eastern deciduous forest, usually under fallen trees or at night foraging in the leaf litter and shaded areas with wet, rocky ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – November. BEHAVIOR: Nocturnal, taking cover during the day under stones, fallen trees, and dead leaves. Adults and larvae eat snails, at least in part, but probably also take other

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kinds of food, such as white grubs of the May (June) Beetle (Phyllophaga spp.). Adults and larvae probably hibernate in fallen trees, in deep fissures in rock outcroppings, or in rodent burrows. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Maronetus) unistriatus (Darlington) 1931 ONE-LINED SNAIL-EATING BEETLE Nomaretus unistriatus Darlington, 1931:149

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Uplands, 1600 meters altitude, in mixed forests, often along slopes. MICROHABITAT: Shaded areas on moderately moist ground rich in leaf litter and humus. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – April, June, and October. BEHAVIOR: Nocturnal, taking cover during the day in leaf litter and humus. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Scaphinotus) vandykei Roeschke 1907 VAN DYKE’S SNAIL-EATING BEETLE Scaphinotus Van Dykei Roeschke, 1907:136 Scaphinotus fuchsi Roeschke, 1907:570

Distribution: Native, New World. USA – AZ. Way of Life: MACROHABITAT: Uplands to mountains, 1830 – 2440 meters altitude, in coniferous forests, often near water bodies. MICROHABITAT: On wet ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – October. BEHAVIOR: Nocturnal, taking cover during the day under stones and bits of fallen trees. References: Erwin (2001), Larochelle & Larivière (2003), Van Dyke (1938), Data from NMNH collection Scaphinotus (Stenocantharis) velutinus (Ménétriés) 1843 VELVET SNAIL-EATING BEETLE Cychrus velutinus Ménétriés, 1843:53 Pemphus longipes Casey, 1897:339 Pemphus opacus Casey, 1899:97

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 1036 meters altitude, in mature coniferous forests (e.g. Sequoia). MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Gidaspow (1973), Larochelle & Larivière (2003), Data from NMNH, UASM collections

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Scaphinotus (Brennus) ventricosus (Dejean) 1831 POTBELLIED SNAIL-EATING BEETLE Cychrus Ventricosus Dejean, 1831:527 Cychrus lativentris Motschulsky, 1850:358 Cychrus ovalis Motschulsky, 1859:162 Cychrus alternatus Motschulsky, 1859:163 Cychrus fuchsianus Rivers, 1890:71 Brennus symmetricus Casey, 1897:319 Brennus strictus Casey, 1897:322 Brennus brevicollis Casey, 1920:178

Distribution: Native, New World. USA – CA, OR. Way of Life: MACROHABITAT: Lowlands to midlands, sea level – 515 meters altitude, in mostly deciduous forests (e.g., Quercus) and poison-oak thickets, as well as on open grassy knolls and edges of reservoirs. MICROHABITAT: Shaded and open ground areas on moderately moist ground, sometimes sandy soils. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. Moderate runner. SEASONAL OCCURRENCE: Adults have been found in January – July, and September – December, teneral adults in late March. BEHAVIOR: Nocturnal, taking cover during the day in fallen trees or under their loose bark. Adults are gregarious. Occasional climber on fallen dead trees while foraging. References: Erwin (2001), Gidaspow (1968), Larochelle & Larivière (2003), Data from NMNH collection Scaphinotus (Scaphinotus) viduus (Dejean) 1826 BEREFT SNAIL-EATING BEETLE Cychrus Viduus Dejean, 1826:12 Cychrus Leonardii T.W. Harris, 1839:193 Cychrus viduus irregularis Beutenmüller, 1903:513

Distribution: Native, New World. CANADA – NB, NS, PQ; USA – CT, DC, DE, GA, MA, ME, MD, NC, NH, NJ, NY, OH, PA, VA, VT, WV. Way of Life: MACROHABITAT: Midlands to uplands, 914 – 2030 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir). At lower altitudes in mixed and deciduous forests (e.g., Quercus spp., Acer spicatum Lam.) and adjacent fields. MICROHABITAT: On shaded wet ground covered with deep leaf litter, usually close to water. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – December, and July – September, teneral adults in July and September. BEHAVIOR: Nocturnal, taking cover during the day under stones, leaf litter, loose bark of fallen trees, under fallen trees, bits of fallen trees, and moss carpets. Adults are active on the surface in rainy weather. Adults have been found over wintering under the loose bark of trees, and in small cells dug under fallen trees, fallen tree-stumps, and pieces of wood. Adult are predaceous on snails, slugs, and caterpillars. Larvae over winter under fallen trees; they are gregarious. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

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Scaphinotus (Steniridia) violaceus carolinae (Valentine) 1935 CAROLINA SNAIL-EATING BEETLE Steniridia violaceus carolinae Valentine, 1935:358

Distribution: Native, New World. USA – NC. Way of Life: MACROHABITAT: Midlands, 670 meters altitude, in deciduous and mixed forests. MICROHABITAT: Ground-dwelling. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Steniridia) violaceus violaceus (LeConte) 1863 VIOLET SNAIL-EATING BEETLE Cychrus violaceus LeConte, 1863:4

Distribution: Native, New World. USA – GA, NC, SC. Way of Life: MACROHABITAT: Midlands, 1340 meters altitude, in deciduous and mixed forests. MICROHABITAT: Shaded areas on moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – July, and September – November. BEHAVIOR: Nocturnal, taking cover during the day at the base of old decaying tree-stumps or under the loose bark of fallen trees. Occasional climber on tree-stumps while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Valentine (1935), Data from NMNH collection Scaphinotus (Scaphinotus) webbi Bell 1959 WEBB’S SNAIL-EATING BEETLE Scaphinotus webbi Bell, 1959:11

Distribution: Native, New World. USA – PA, VA, WV. Way of Life: MACROHABITAT: Lowlands, 285 meters altitude, in deciduous forests (e.g., Quercus), often along brooks and near springs. MICROHABITAT: Shaded areas on moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – August, teneral adults in July. BEHAVIOR: Nocturnal, taking cover during the day under large stones. Adult are predaceous on snails. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH, UASM collections Sphaeroderus Dejean 1826 FALSE SNAIL-EATING BEETLES (Plate 21) Sphaeroderus Dejean, 1826:2

Number of worldwide species: 5. Number of Western Hemisphere species: 5. Taxonomy: Stable. Adelphotaxon: Scaphinotus Dejean 1826

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Type species: Cychrus stenostomus Weber, 1801 Distribution: Nearctic. Habitat: According to Lindroth (1961), species in this genus hibernate as adults, unlike most other Cychrini. They are inhabitants of mesic forests. References: Erwin (2001), Lindroth (1961) Sphaeroderus bicarinatus (LeConte) 1853 TWO-RIDGED FALSE SNAIL-EATING BEETLE Cychrus bicarinatus LeConte, 1853:399 Sphaeroderus multicarinatus Darlington, 1931:151

Distribution: Native, New World. USA – AL, GA, NC, SC, TN. Way of Life: MACROHABITAT: Midlands to uplands, 900 – 2030 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir, (at lower altitudes in mixed and deciduous forests, e.g., Quercus spp., Acer spicatum Lam.). MICROHABITAT: Shaded areas on moderately moist ground covered with deep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – October, and December, teneral adults in late August. BEHAVIOR: Nocturnal, taking cover during the day in leaf litter, under the loose bark of fallen trees, in and under rotten fallen trees, under stones, and under moss carpets. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Sphaeroderus canadensis canadensis Chaudoir 1861 CANADIAN FALSE SNAIL-EATING BEETLE Sphaeroderus canadensis Chaudoir, 1861:498 Sphaeroderus palpalis Motschulsky, 1866:312 Sphaeroderus canadensis Blanchardi Leng, 1916:41

Distribution: Native, New World. CANADA – NB, NS, ON, PQ; USA – CT, KY, MA, MD, ME, MI, NC, NH, NY, OH, PA, SC, TN, VA, VT, WV. Way of Life: MACROHABITAT: Lowlands, sea level – 457 meters altitude, in coniferous, deciduous, and mixed forests along rivers, brooks, the seashore, and on slopes. MICROHABITAT: Shaded areas on moderately moist (mostly) or wet ground with deep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – November, copulating pairs in April – July, gravid females in late May to late July, teneral adults in May, and July – September. BEHAVIOR: Mostly Nocturnal, taking cover during the day in leaf litter, under fallen trees, stones, and moss carpets. Larval habitat similar to adult, in leaf litter. Coitus lasting 101 – 142 minutes. Adults have been found over wintering on higher and drier ground, at the summit of wooded slopes and hillocks in leaf litter. Adult are predaceous on snails, lepidopterous caterpillars and pupae. Occasional climber on tree trunks while foraging. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Notes: Gidaspow (1973) regarded this name as in the genus Nomaretus.

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Sphaeroderus canadensis lengi Darlington 1933 LENG’S FALSE SNAIL-EATING BEETLE Sphaeroderus canadensis lengi Darlington, 1933:63

Distribution: Native, New World. USA – GA, NC, SC, TN, WV. Way of Life: MACROHABITAT: Midlands to uplands, 915 – 2030 meters altitude, in coniferous forests (e.g., Picea rubens Sarg., Abies fraseri (Pursh) Poir, (at lower altitudes in mixed and deciduous forests, e.g., Quercus spp., Acer spicatum Lam.). MICROHABITAT: Shaded areas on moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in March – November. BEHAVIOR: Nocturnal, taking cover during the day beneath fallen trees and in leaf litter. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Sphaeroderus indianae (Blatchley) 1910 INDIANA FALSE SNAIL-EATING BEETLE Cychrus stenostomus indianae Blatchley, 1910:42

Distribution: Native, New World. USA – IN, KY, OH, TN, VA, WV. Way of Life: MACROHABITAT: Lowlands, 182 – 377 meters altitude, in forests. MICROHABITAT: Shaded areas on moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February, April, and July – October. BEHAVIOR: Adults found over wintering in leaf litter. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Sphaeroderus nitidicollis brevoorti LeConte 1848 BREVOORT’S FALSE SNAIL-EATING BEETLE Sphaeroderus Brevoorti LeConte, 1848:343 Sphaeroderus granulosus Chaudoir, 1861:497

Distribution: Native, New World. CANADA – MB, NB, NS, ON, PQ, SK; USA – ME, MN, NH, NY, VT. Way of Life: MACROHABITAT: Lowlands to uplands, and subalpine zone, sea level – 1600 meters altitude, in coniferous and mixed forests or at their edges. Adults occur along rivers, brooks, at the seashore, and on slopes. MICROHABITAT: Shaded area on wet or moist ground covered with deep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – November; copulating pairs seen in June – November; gravid females in spring, and August – November; teneral adults in August and the autumn. BEHAVIOR: Mostly nocturnal, taking cover during the day in leaf litter, under fallen trees or stones; sometimes adults are active on cloudy days. Occasional climber on tree trunks while foraging. Larval habits are similar to adults. According to Larochelle & Larivière (2003), “adults have been found over wintering on higher and drier ground, in woods; under moss and in leaf litter, at the base of tree-stumps and trees, under the loose bark of fallen trees, in fallen rotten branches, and under stones.” Adult and larvae are predaceous on snails. Coitus has been recorded as lasting up to 72 hours. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection

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Sphaeroderus nitidicollis nitidicollis Guérin-Méneville 1829 POLISHED-COLLAR FALSE SNAIL-EATING BEETLE Sphaeroderus nitidicollis Guérin-Méneville, 1829: pl. 7

Distribution: Native, New World. CANADA – NF, PQ. Way of Life: MACROHABITAT: Lowlands and subalpine zone, sea level – 51 meters altitude, in deciduous, mixed, or coniferous forests, and in thickets. Adults occur along rivers, brooks, at the seashore. MICROHABITAT: Shaded areas on wet or moist ground covered with deep leaf litter. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – September; copulating pairs seen in July; teneral adults in July – August. BEHAVIOR: Nocturnal, taking cover during the day under dead leaves or in moss, fallen trees, or under stones. Adult are predaceous on snails. Occasional climber on tree trunks while foraging. Coitus has been recorded as lasting up to 22 hours. Adults hibernate. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Sphaeroderus nitidicollis schaumii Chaudoir 1861 SCHAUM’S FALSE SNAIL-EATING BEETLE Sphaeroderus Schaumii Chaudoir, 1861:499

Distribution: Native, New World. USA – IL, MI, OH, VA, WV. Way of Life: MACROHABITAT: Lowlands to midlands, 280 – 1100 meters altitude, in deciduous forests (e.g., Acer; Celtis). MICROHABITAT: Shaded areas on moderately moist ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in June – November, teneral adults in August. BEHAVIOR: Nocturnal, taking cover during the day under dead leaves, fallen trees and wood bits, or under stones. Adult are predaceous on snails. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Sphaeroderus stenostomus lecontei Dejean 1826 LECONTE’S FALSE SNAIL-EATING BEETLE Sphaeroderus Lecontei Dejean, 1826:15 Sphaeroderus niagarensis Laporte de Castelnau, 1833:390 Sphaeroderus lecontei diffractus Casey, 1914:25

Distribution: Native, New World. CANADA – MB, NB, NF, NS, ON, PE, PQ, SK; USA – AL, CT, FL, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, MS, NC, NH, NJ, NY, OH, PA, RI, SC, TN, VA, VT, WI, WV. Way of Life: MACROHABITAT: Lowlands to midlands, and subalpine zone, sea level – 1067 meters altitude. Eurytopic, in mixed, deciduous, and coniferous forests often along rivers, brooks, lakes, and at the seashore. Adults are also sometimes found in forest clearings and their edges, along adjoining roadsides, and in sand pits, gravel pits, fields, swamps, and marshes. MICROHABITAT: Shaded or half-shaded areas on moist or wet ground covered with deep leaf litter, especially in dense hard-wood forests with Cornus, Viburnum, Acer, Alnus, Sorbus, & co. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running.

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SEASONAL OCCURRENCE: Adults have been found in January – December; copulating pairs in April – July, September – November; ovipositing females in May – July; late instar larvae in August; teneral adults in July – September. BEHAVIOR: Mostly nocturnal, taking cover during the day in leaf litter, under fallen trees and stones, in mosses, or under the loose bark of fallen trees; may invade river banks and other open sites at night; sometimes active on cloudy days. Adults are gregarious in winter. Larvae found in leaf litter. Adults found over wintering on higher and drier ground, in woods and at their edges, in leaf litter, in rotten tree-stumps and fallen trees, under clods of earth, and under stones. Adult are predaceous on snails, slugs, and lepidopterous caterpillars. Occasional climber while foraging. Coitus has been recorded as lasting up to 72 hours. References: Erwin (2001), Larochelle & Larivière (2003), Lindroth (1955), Data from NMNH collection Sphaeroderus stenostomus stenostomus (Weber) 1801 NARROW-MOUTHED FALSE SNAIL-EATING BEETLE Cychrus stenostomus Weber, 1801:43 Sphaeroderus stenostomus aequalis Casey, 1920:172 [New synonymy]

Distribution: Native, New World. USA – DC, DE, MD, NC, NJ, NY, PA, VA, WV. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 1122 meters altitude, in deciduous and coniferous forests and at the borders of eutrophic marshes. MICROHABITAT: Shaded areas on dry ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – November; copulating pairs in April, October – November; teneral adults in July. BEHAVIOR: Mostly Nocturnal, taking cover during the day under dead leaves, stones, and fallen trees, sometimes active on cloudy days. Adults hibernate. Adult are predaceous on snails and slugs. References: Erwin (2001), Larochelle & Larivière (2003), Data from NMNH collection Notes: Casey’s single type specimen from PA (NMNH) matches perfectly one of three specimens collected together on the Blue Ridge summit in Franklin County. The only differences between Casey’ specimen and the matching one and the other two specimens are that the pronotal base is not extensively punctulate and the elytra are a bit more parallel-sided. The other two specimens are perfectly good S. s. stenomus (Weber). In a series of 298 specimens in NMNH from throughout the range of S. s. stenomus (Weber), both these attributes are variable. Since the variation exists in co-collected specimens, subspecies status is not possible. CICINDELITAE Collyridini Brullé 1834 COMB-MOUTHED BEETLES Collyridini Brullé, 1834:27

Number of worldwide genera: One. Number of Western Hemisphere genera: One. Taxonomy: Stable. Adelphotaxon: Pogonostoma Klug 1835 Geographic status: Neotropical. References: Erwin (2001), Lorenz (2005), Naviaux (1998)

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Ctenostoma Klug 1821 COMB-MOUTHED BEETLES (Figure 22) Ctenostoma Klug, 1821:302 Caris, Fisher von Waldheim, 1821:99 Procephalus Laporte de Castelnau, 1834:35 [Bousquet (2002:42) demonstrated, followed by Lorenz (2005:153), that this name is a junior subjective synonym of the nominotypical subgenus of Ctenostoma and that a new name is needed for this subgenus of species. Naviauxiana Erwin, new replacement name for Procephalus Laporte de Castelnau, 1834 [valid subgenus for native Neotropical species] Myrmecilla Lacordaire, 1843:93 [valid subgenus for native Neotropical species] Neoprocephalus Naviaux, 1998:59 [valid subgenus for native Neotropical species] Microprocephalus Naviaux, 1998:84 [valid subgenus for native Neotropical species] Euctenostoma Naviaux, 1998:105 [valid subgenus for native Neotropical species] Paractenosoma Naviaux, 1998:170 [valid subgenus for native Neotropical species] Salvioides Naviaux, 1998:174 [valid subgenus for native Neotropical species]

Number of worldwide species: 109. Number of Western Hemisphere species: 109. Taxonomy: Stable. Adelphotaxon: Pogonostoma Klug 1835 Type species: “Ctenostoma formacarium F.” [non Fabricius 1801], = Ctenostoma jekelii Chevrolat, 1858. Distribution: México, south to Paraguay and Bolivia. Habitat: Arboreal: both canopy and understory of rainforests and tropical moist forests of the Mata Atlântica; some altitude records associated with the Andes and Central America indicate they also occur in cloud forests. These long-legged beetles, many of which are ant-shaped and brachypterous, are found on twigs and foliage in the canopy and understory of moist tropical forests, rain forests, and likely cloud forests; also adults have been found in fogging samples of the tall river-side grass, Gynerium sagittatum (Aubl.) Beauv and stands of the bamboo, Guadua weberbaueri Pilger. Larvae use holes in rotting twigs and perhaps bamboo culms much as other tiger beetle larvae use holes in the ground. Some species make their burrows in the end of broken twigs, others in the lateral surfaces of the twig. The burrows are enlarged as the larva grows. Both adults and larvae are predatory on insects. Although Zikan (1929) reported observations on the way of life of some species in South America, particularly Brazil, the genus remains inadequately known. Naviaux (1998) noted that adults of some species seem to mimic cerambycid beetles; perhaps it is the other way round since all carabids possess well-developed chemical defense systems (pygidial glands and deliverables). References: Erwin (2001), Cassola (2001), Erwin (1991), Naviaux (1998, 2002), Pearson et al. (1999a), Pearson et al. (1999b), Zikan (1929) Ctenostoma (Myrmecilla) abbreviatum Naviaux 1998 SHORT COMB-MOUTHED BEETLE Ctenostoma abbreviatum Naviaux, 1998:143

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence

SPECIES ACCOUNTS / CARABIDAE

flightless thus vagility limited to walking or running. SEASONAL BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

OCCURRENCE:

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173

Unknown.

Ctenostoma (Ctenostoma) acciavatti Naviaux 1998 ACCIAVATTI’S COMB-MOUTHED BEETLE Ctenostoma acciavatti Naviaux, 1998:91

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: The adult holotype was found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) aeneum Naviaux 1998 BRASSY COMB-MOUTHED BEETLE Ctenostoma aeneum Naviaux, 1998:42

Distribution: Native, New World. COSTA RICA; NICARAGUA; PANAMÁ. Way of Life: MACROHABITAT: Lowlands to midlands, 300 – 1450 meters altitude, in forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. One adult found in association with the plant, Ocotea whitei Woodson. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) agnatum agnatum Chaudoir 1860 SMALL-JAWED COMB-MOUTHED BEETLE Ctenostoma agnatum Chaudoir, 1860:280

Distribution: Native, New World. BRAZIL, ECUADOR, PERÚ. Way of Life: MACROHABITAT: Lowlands to mountains, 100 – 2228 meters altitude, in rainforest and cloud forests. MICROHABITAT: Canopy, closed, at an average of 37 meters height. Also, in suspended dry leaves in the canopy and on leaves of undergrowth in primary forest. DISPERSAL ABILITIES: Brachypterous, flight wing less that length of elytra, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February – March, June, and September – October. BEHAVIOR: See description under genus, above. Many examples were obtained from canopy foggings in the Yasuni area of eastern Ecuador, in wet, dry and transitional seasons, thus members of this species are active throughout the year. Adults of this species have been observed “playing dead” when disturbed. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Pearson et al. (1999a), Data from D. Pearson, NMNH collections

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Ctenostoma (Myrmecilla) agnatum fuscum Naviaux 1998 SOMBER COMB-MOUTHED BEETLE Ctenostoma agnatum fuscum Naviaux, 1998:164

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 350 meters altitude, in rainforest with bamboo patches. MICROHABITAT: Suspended dry leaves in bamboo stand (Guadua weberbaueri Pilger). DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: The adult holotype was found in October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) albofasciatum Chaudoir 1850 WHITE-LINED COMB-MOUTHED BEETLE Ctenostoma albofasciatum Chaudoir, 1850:16

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) angustoobliquatum W. Horn 1925 NARROWLY-OBLIQUE COMB-MOUTHED BEETLE Ctenostoma angusto-obliquatum W. Horn, 1925:132

Distribution: Native, New World. COSTA RICA. Way of Life: MACROHABITAT: Lowlands to uplands, 75 – 1650 meters altitude, in forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – June, and August. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Naviauxiana) arnaudi Naviaux 1998 ARNAUD’S COMB-MOUTHED BEETLE Ctenostoma arnaudi Naviaux, 1998:37

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands to mountains, 650 – 2228 meters altitude, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – February, May, July and September. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005)

SPECIES ACCOUNTS / CARABIDAE

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175

Note: Pearson et al. (1999a) cast doubt on the western slope 650 meter altitude record in Esmeraldas Province believing this species is an Amazon Basin one, thus not occurring in the west. Ctenostoma (Myrmecilla) asperulum Bates 1868 ROUGH COMB-MOUTHED BEETLE Ctenostoma asperulum Bates, 1868:278

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) bahiaense Naviaux 1998 BAHIA COMB-MOUTHED BEETLE Ctenostoma bahiaensis Naviaux, 1998:97

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES : Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) batesii Chaudoir 1860 BATES’ COMB-MOUTHED BEETLE Ctenostoma Batesii Chaudoir, 1860:276

Distribution: Native, New World. BRAZIL; ECUADOR. Way of Life: MACROHABITAT: Lowlands, 216 – 250 meters altitude, in rainforest. MICROHABITAT: Canopy, closed, at an average of 37 meters height. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February, June, and October. BEHAVIOR: See description under genus, above. Many examples were obtained from canopy foggings in the Yasuni area of eastern Ecuador, in wet, dry and transitional seasons, thus members of this species are active throughout the year. References: Erwin (2001, field notes), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) bicristatum Chaudoir 1860 DOUBLE-CRESTED COMB-MOUTHED BEETLE Ctenostoma bicristatum Chaudoir, 1860:282

Distribution: Native, New World. BRAZIL.

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Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) bifasciatum bifasciatum Dejean 1831 DOUBLE-MARKED COMB-MOUTHED BEETLE Ctenostoma Bifasciatum Dejean, 1831:272

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) bifasciatum horni Naviaux 1998 HORN’S COMB-MOUTHED BEETLE Ctenostoma bifasciatum horni Naviaux, 1998:112

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) bondari W. Horn 1939 BONDAR’S COMB-MOUTHED BEETLE Ctenostoma Bondari W. Horn, 1939:38

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) brendelli Naviaux 1998 BRENDELL’S COMB-MOUTHED BEETLE Ctenostoma brendelli Naviaux, 1998:80

Distribution: Native, New World. COSTA RICA; PANAMÁ.

SPECIES ACCOUNTS / CARABIDAE

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177

Way of Life: MACROHABITAT: Lowlands, 133 meters altitude, in dry forest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Myrmecilla) brevicorne W. Horn 1898 SHORT-HORNED COMB-MOUTHED BEETLE Ctenostoma brevicorne W. Horn, 1898:29

Distribution: Native, New World. VENEZUELA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Adults are found on twigs and foliage in the understory of moist tropical forest types, and larvae are found in burrows in twigs and small branches. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from D. Pearson collection Ctenostoma (Microprocephalus) brevilabre W. Horn 1931 SHORT-LIPPED’S COMB-MOUTHED BEETLE Ctenostoma brevilabre W. Horn, 1931:101

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) breviusculum von Mannerheim 1837 MANNERHEIM’S COMB-MOUTHED BEETLE Ctenostoma breviusculum von Mannerheim, 1837:20

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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Ctenostoma (Neoprocephalus) brunneum Naviaux 1998 BROWNISH COMB-MOUTHED BEETLE Ctenostoma brunneum Naviaux, 1998:66

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands, 650 meters altitude, in rain forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Neoprocephalus) cassolai Naviaux 1998 CASSOLA’S COMB-MOUTHED BEETLE Ctenostoma cassolai Naviaux, 1998:67

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands, 950 meters altitude, in cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: The adult holotype was found in March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) cayennense Naviaux 1998 CAYENNE COMB-MOUTHED BEETLE Ctenostoma cayennensis Naviaux, 1998:53

Distribution: Native, New World. FRENCH GUIANA. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February, April, August, October – November, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Ctenostoma) chaudoiri (W. Horn) 1895 CHAUDOIR’S COMB-MOUTHED BEETLE Procephalus Chaudoiri W. Horn, 1895:353 Ctenostoma Chaudoiri Gounelli W. Horn, 1898:27 Ctenostoma chaudoiri gounellei W. Horn, 1926:94

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Alantic forest (Mata Atlântica). The type locality of “gounelli” was Mata Atlântica when the type specimen was collected. At present, that part of the Tropical Moist Forest of Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably

SPECIES ACCOUNTS / CARABIDAE

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179

capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – April. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from CAS collection Ctenostoma (Myrmecilla) compactum Naviaux 1998 COMPACT COMB-MOUTHED BEETLE Ctenostoma compactum Naviaux, 1998:167

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 350 – 450 meters altitude, in rainforest. MICROHABITAT: Canopy of Astrocaryum palm, in the dry suspended fronds skirting the trunk. DISPERSAL ABILITIES: Brachypterous, flight wing less that length of elytra, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August, and October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) coracinum Naviaux 1998 BLACKISH COMB-MOUTHED BEETLE Ctenostoma coracinum Naviaux, 1998:141

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Midlands to uplands, 1460 – 1600 meters altitude, in forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February, and October – November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Paractenosoma) corculum Bates 1868 LITTLE-HEART COMB-MOUTHED BEETLE Ctenostoma corculum Bates, 1868:277

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) crucifrons W. Horn 1911 CROSS-FACED COMB-MOUTHED BEETLE Ctenostoma crucifrons W. Horn, 1911:154

Distribution: Native, New World. PERÚ.

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Way of Life: MACROHABITAT: Lowlands to midlands, 450 – 705 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – August, and October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from CAS collection Ctenostoma (Neoprocephalus) cylindratum Naviaux 1998 ROLLER-TOOL COMB-MOUTHED BEETLE Ctenostoma cylindratum Naviaux, 1998:64

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Lowlands, 200 – 400 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in October – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) davidsoni Naviaux 1998 DAVIDSON’S COMB-MOUTHED BEETLE Ctenostoma davidsoni Naviaux, 1998:62

Distribution: Native, New World. COSTA RICA. Way of Life: MACROHABITAT: Lowlands to midlands, 75 – 610 meters altitude, in rainforest and tropical dry forest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Myrmecilla) dentifrons W. Horn 1901 TOOTH-FACED COMB-MOUTHED BEETLE Ctenostoma dentifrons W. Horn, 1901:353

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). When the type was collected in the state of Espirito Santo, Brazil, the Mata Atlântica in Brazil was still growing. At present, nearly all the forests of this Brazilian state have been converted to non-forest vegetation. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

SPECIES ACCOUNTS / CARABIDAE

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181

Ctenostoma (Neoprocephalus) deuvei Naviaux 1998 DEUVE’S COMB-MOUTHED BEETLE Ctenostoma deuvei Naviaux, 1998:70

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 300 – 450 meters altitude, in rainforest. MICROHABITAT: Canopy of Caña Brava grasses [Gynerium sagittatum (Aubl.) Beauv.] on river margin. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – August, and October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) dokhturowi W. Horn 1898 DOKHTUROW’S COMB-MOUTHED BEETLE Ctenostoma Dokhturowi W. Horn, 1898:32

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) dormeri W. Horn 1898 DORMER’S COMB-MOUTHED BEETLE Ctenostoma Dormeri W. Horn, 1898:196

Distribution: Native, New World. COLOMBIA; ECUADOR. Way of Life: MACROHABITAT: Midlands to mountains, 900 – 2200 meters, in upper rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June – July, and November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from D. Pearson collection Ctenostoma (Naviauxiana) durantoni Naviaux 1998 DURANTON’S COMB-MOUTHED BEETLE Ctenostoma durantoni Naviaux, 1998:58

Distribution: Native, New World. BRAZIL; FRENCH GUIANA. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February – July, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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Ctenostoma (Naviauxiana) ebenina Bates 1868 EBONY COMB-MOUTHED BEETLE Ctenostoma ebeninum Bates, 1868:277

Distribution: Native, New World. BRAZIL; BOLIVIA; FRENCH GUIANA; VENEZUELA. Way of Life: MACROHABITAT: Lowlands to midlands, 200 – 800 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April, and October – November. BEHAVIOR: See description under genus, above. Adults are attracted to lights at night. References: Erwin (2001), Naviaux (1998), Pearson, et al. (1999b), Data from D. Pearson collection Ctenostoma (Euctenostoma) eburatum Bates 1872 IVORY COMB-MOUTHED BEETLE Ctenostoma eburatum Bates, 1872:49

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Lowlands, 50 – 100 meters altitude in lowland rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) ecuadorense Naviaux 1998 ECUADORIAN COMB-MOUTHED BEETLE Ctenostoma ecuadorensis Naviaux, 1998:46

Distribution: Native, New World. COLOMBIA; ECUADOR. Way of Life: MACROHABITAT: Lowlands to uplands, 70 – 1800 meters altitude, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – March, and October – November. BEHAVIOR: See description under genus, above. Adults are attracted to lights. References: Erwin (2001), Naviaux (1998), Pearson et al. (1999a) Ctenostoma (Naviauxiana) erwini Naviaux 1998 ERWIN’S COMB-MOUTHED BEETLE Ctenostoma erwini Naviaux, 1998:48

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 850 meters altitude, in rainforest and cloud forests. MICROHABITAT: Adults are found on twigs and foliage in the shady understory of moist tropical forest types, and larvae are found in burrows in twigs and small

SPECIES ACCOUNTS / CARABIDAE

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183

branches. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: Adults are predaceous on small arboreal arthropods, seeking them on twigs and leaves of forest trees and shrubs. Adults occasionally found at lights at night. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from NMNH collection Ctenostoma (Euctenostoma) ferum Naviaux 2005 WILD COMB-MOUTHED BEETLE Ctenostoma ferum Naviaux, 2005:267

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Naviaux (2005) Ctenostoma (Myrmecilla) flexuosum Naviaux 1998 FLEXED-FASCIA COMB-MOUTHED BEETLE Ctenostoma flexuosum Naviaux, 1998:133

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) formicarium (Fabricius) 1801 ANT-LIKE COMB-MOUTHED BEETLE Collyris formicarium Fabricius, 1801:226 Ctenostoma Jacquieri Dejean, 1831:271 Ctenostoma formicarium rectofasciata W. Horn, 1913:01

Distribution: Native, New World. BRAZIL; FRENCH GUIANA; PERÚ. Way of Life: MACROHABITAT: Lowlands, 32 – 203 meters altitude in lowland rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from CAS collection

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Ctenostoma (Euctenostoma) fryi Chaudoir 1865 FRY’S COMB-MOUTHED BEETLE Ctenostoma Fryi Chaudoir, 1865:49

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) gautardi Chaudoir 1869 GAUTARD’S COMB-MOUTHED BEETLE Ctenostoma Gautardi Chaudoir, 1869:22

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) germaini W. Horn 1902 GERMAIN’S COMB-MOUTHED BEETLE Ctenostoma Germaini W. Horn, 1902:372

Distribution: Native, New World. BOLIVIA. Way of Life: MACROHABITAT: Uplands, 2000+ meters altitude, in montane forests, Yungas. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Pearson, et al. (1999b) Ctenostoma (Myrmecilla) globifrons W. Horn 1898 SPHERICAL-FACED COMB-MOUTHED BEETLE Ctenostoma globifrons W. Horn, 1898:31

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Alantic forest (Mata Atlântica). The type locality of “globifrons” was Mata Atlântica when the type specimen was collected. At present, that part of the tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in May – June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

SPECIES ACCOUNTS / CARABIDAE

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185

Ctenostoma (Neoprocephalus) guatemalense Brouerius van Nidek 1960 GUATEMALAN COMB-MOUTHED BEETLE Ctenostoma guatemalense Brouerius van Nidek, 1960:8

Distribution: Native, New World. GUATEMALA. Way of Life: MACROHABITAT: Midlands, 1250 meters altitude, in lower montane rain forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) guyanense W. Horn 1998 GUIANAN COMB-MOUTHED BEETLE Ctenostoma guyanense Naviaux, 1998:90

Distribution: Native, New World. FRENCH GUIANA. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – February, September – October, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) heydeni W. Horn 1894 HEYDEN’S COMB-MOUTHED BEETLE Ctenostoma Heydeni W. Horn, 1894:109

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) hirsutum W. Horn 1892 HAIRY COMB-MOUTHED BEETLE Ctenostoma hirsutum W. Horn, 1892:46 Ctenostoma macilentum Klug, 1898:29.

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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Ctenostoma (Neoprocephalus) hovorei Naviaux & Brzoska 2005 PACO’S COMB-MOUTHED BEETLE Ctenostoma hovorei Naviaux & Brzoska, 2005:24

Distribution: Native, New World. COSTA RICA. Way of Life: MACROHABITAT: Midlands, 1350 meters altitude, in cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – July. BEHAVIOR: See description under genus, above. References: Naviaux & Brzoska (2005) Ctenostoma (Neoprocephalus) ibidion ibidion Dohrn 1880 DOHRN’S COMB-MOUTHED BEETLE Ctenostoma ibidion Dohrn, 1880:372

Distribution: Native, New World. COSTA RICA; VENEZUELA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) ibidion jolyi Naviaux 1998 JOLY’S COMB-MOUTHED BEETLE Ctenostoma ibidion jolyi Naviaux, 1998:73

Distribution: Native, New World. ECUADOR, VENEZUELA. Way of Life: MACROHABITAT: Uplands to mountains, 1600 – 2300 meters altitude, in cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – July, and October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Myrmecilla) ichneumoneum ellipticum Naviaux 1998 ELLIPSUS-MARKED COMB-MOUTHED BEETLE Ctenostoma ichneumoneum ellipticum Naviaux, 1998:140

Distribution: Native, New World. BRAZIL; PARAGUAY. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March, and November – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998, 2002)

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187

Ctenostoma (Myrmecilla) ichneumoneum ichneumoneum Dejean 1826 WASP-LIKE COMB-MOUTHED BEETLE Ctenostoma Ichneumoneum Dejean, 1826:436 Ctenostoma ichneumoneum interruptum Chaudoir, 1860:283

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – February and November – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) immaculatum W. Horn 1925 NON-SPOTTED COMB-MOUTHED BEETLE Ctenostoma immaculatum W. Horn, 1925:182

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands, 650 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) inca Naviaux 1998 INCA COMB-MOUTHED BEETLE Ctenostoma inca Naviaux, 1998:122

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands to mountains, 350 – 2228 meters altitude, in rainforest and cloud forests. MICROHABITAT: Dry hanging fronds of Astrocaryum palm in the upper floodplain forest on the Rio Manu. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in September – October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from NMNH collection Ctenostoma (Myrmecilla) infimum Naviaux 1998 LOWLY COMB-MOUTHED BEETLE Ctenostoma infimum Naviaux, 1998:156

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence

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flightless thus vagility limited to walking or running. SEASONAL BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998, 2002)

OCCURRENCE:

Unknown.

Ctenostoma (Naviauxiana) insigne Chaudoir 1860 MARKED COMB-MOUTHED BEETLE Ctenostoma insigne Chaudoir, 1860:274

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Lowlands, 100 – 300 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998, 2002) Ctenostoma (Neoprocephalus) intermedium Naviaux 1998 INTERMEDIATE COMB-MOUTHED BEETLE Ctenostoma intermedium Naviaux, 1998:71

Distribution: Native, New World. VENEZUELA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) jekelii jekelii Chevrolat 1858 JEKEL’S COMB-MOUTHED BEETLE Ctenostoma formicarum, [sic] Fabricius: Klug, 1821:304 Ctenostoma Jekelii Chevrolat, 1858:484 Ctenostoma Jekeli W. Horn, 1898:28

Distribution: Native, New World. BRAZIL; FRENCH GUIANA. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Adults are found on twigs and foliage in the shady understory of moist tropical forest types, and larvae are found in burrows in twigs and small branches. DISPERSAL ABILITIES: Brachypterous, wing length variable, however adults are flightless, thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January. BEHAVIOR: Adults are predaceous on small arboreal arthropods, seeking them on twigs and leaves of forest trees and shrubs. References: Erwin (2001), Naviaux (1998), Data from D. Pearson collection

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189

Ctenostoma (Ctenostoma) jekelii raresculptum W. Horn 1898 RARE-SCULPTURED COMB-MOUTHED BEETLE Ctenostoma Jekeli raresculpta W. Horn, 1898:28

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Alantic forest (Mata Atlântica). The type locality of “raresculptum” was Mata Atlântica when the type specimen was collected. At present, that part of the tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, wing length variable, however adults are flightless, thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) johnsoni Naviaux 1998 JOHNSON’S COMB-MOUTHED BEETLE Ctenostoma intermedium Naviaux, 1998:83

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands, 500 meters altitude, in wet forest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) klugeanum W. Horn 1915 KLUG’S COMB-MOUTHED BEETLE Ctenostoma Klugeanum W. Horn, 1915:424 Ctenostoma Klugi W. Horn, 1898:28

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) laeticolor Bates 1878 JOYFUL-COLORED COMB-MOUTHED BEETLE Ctenostoma laeticolor Bates, 1878:588

Distribution: Native, New World. COSTA RICA; NICARAGUA; PANAMÁ. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably

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capable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) landolti Steinheil 1877 LANDOLT’S COMB-MOUTHED BEETLE Ctenostoma Landolti Steinheil, 1877:48

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) longipalpe Naviaux 1998 LONG-PALPI COMB-MOUTHED BEETLE Ctenostoma longipalpe Naviaux, 1998:40

Distribution: Native, New World. COSTA RICA; PANAMÁ. Way of Life: MACROHABITAT: Lowlands to uplands, 75 – 1550 meters, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – August, and October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from D. Pearson, NMNH collections Ctenostoma (Euctenostoma) luctuosum Chaudoir 1860 MOURNFUL COMB-MOUTHED BEETLE Ctenostoma luctuosum Chaudoir, 1860:277

Distribution: Native, New World. BRAZIL; FRENCH GUIANA, PERÚ. Way of Life: MACROHABITAT: Lowlands, 290 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. One example was obtained from canopy foggings in southeastern Perú, in wet to dry transitional season. Adults are attracted to lights. References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Data from D. Pearson, NMNH collections

SPECIES ACCOUNTS / CARABIDAE

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191

Ctenostoma (Myrmecilla) luteum Naviaux 1998 YELLOWISH COMB-MOUTHED BEETLE Ctenostoma luteum Naviaux, 1998:166

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 350 meters altitude, in rainforest. MICROHABITAT: Suspended dry leaves. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: The adult holotype was found in October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Erwin (field notes), Naviaux (1998), Data from NMNH collection Ctenostoma (Myrmecilla) macilentum Klug 1834 THIN COMB-MOUTHED BEETLE Ctenostoma macilentum Klug, 1834:42

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) maculicorne (Chevrolat) 1856 SPOTTED-HORN COMB-MOUTHED BEETLE Procephalus maculicornis Chevrolat, 1856:352 Ctenostoma sigma Bates, 1881:15

Distribution: Native, New World. BELIZE; COLOMBIA; COSTA RICA; GUATEMALA; MÉXICO – OA, VC, YC; NICARAGUA; PANAMÁ. Way of Life: MACROHABITAT: Lowlands to midlands, 30 – 1000 meters altitude, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – July, and October – November. BEHAVIOR: See description under genus, above. Adults have been captured in a malaise trap and they are attracted to lights at night. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from D. Pearson, CAS, NMNH collections Ctenostoma (Naviauxiana) maculosum Naviaux 1998 SMALL-SPOT COMB-MOUTHED BEETLE Ctenostoma maculosum Naviaux, 1998:47

Distribution: Native, New World. COLOMBIA. Way of Life: MACROHABITAT: Uplands, 2000 meters altitude, in cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable

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of flight. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description under genus, above. Adults occasionally found at lights at night. References: Erwin (2001), Naviaux (1998), Data from NMNH collection Ctenostoma (Euctenostoma) magnum Naviaux 1998 VERY LARGE COMB-MOUTHED BEETLE Ctenostoma magnum Naviaux, 1998:124

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 350 – 450 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – August. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) metallicum (Laporte de Castelnau) 1834 METALLIC COMB-MOUTHED BEETLE Procephalus metallicum Laporte de Castelnau, 1834:36

Distribution: Native, New World. FRENCH GUIANA; GUYANA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) minusculum Naviaux 2002 VERY SMALL COMB-MOUTHED BEETLE Ctenostoma minusculum Naviaux, 2002:266

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in November. BEHAVIOR: See description under genus, above. References: Naviaux (1998) Ctenostoma (Myrmecilla) modicum Naviaux 1998 MODEST COMB-MOUTHED BEETLE Ctenostoma modicum Naviaux, 1998:168

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica), 1100 meters altitude. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES:

SPECIES ACCOUNTS / CARABIDAE

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193

Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) monnei Naviaux 2002 MONNE’S COMB-MOUTHED BEETLE Ctenostoma monnei Naviaux, 2002:265

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Naviaux (2002) Ctenostoma (Naviauxiana) nigrum lecourti Naviaux 1998 LECOURT’S COMB-MOUTHED BEETLE Ctenostoma nigrum lecourti Naviaux, 1998:35

Distribution: Native, New World. BOLIVIA; PERÚ. Way of Life: MACROHABITAT: Lowlands to upland, 260 – 1500 meters altitude, in rainforest and cloud forests, Yungas. MICROHABITAT: Suspended dry leaves. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March – April, and October – November. BEHAVIOR: See description under genus, above. An adult was found in suspended dry leaves in a stand of bamboo (Guadua weberbaueri Pilger) about 2.5 meters off the ground using insecticidal fogging techniques. References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Data from NMNH collection Ctenostoma (Naviauxiana) nigrum nigrum Chaudoir 1860 BLACK COMB-MOUTHED BEETLE Ctenostoma nigrum Chaudoir, 1860:275

Distribution: Native, New World. BRAZIL; COLOMBIA; PERÚ. Way of Life: MACROHABITAT: Lowlands, 350 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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Ctenostoma (Neoprocephalus) nitidum Naviaux 1998 BRILLIANT COMB-MOUTHED BEETLE Ctenostoma nitidum Naviaux, 1998:74

Distribution: Native, New World. BOLIVIA; PERÚ. Way of Life: MACROHABITAT: Lowlands to uplands, 400 – 1500 meters altitude, in rainforest and cloudforests, Yungas. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February, October – December. BEHAVIOR: See description under genus, above. An adult male was found at 2.5 meters off the ground in suspended dry leaves in a stand of bamboo (Guadua weberbaueri Pilger) using insecticidal fogging techniques. References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b) Data from NMNH collection Ctenostoma (Myrmecilla) obliquatum Chaudoir 1860 SLOPED COMB-MOUTHED BEETLE Ctenostoma obliquatum Chaudoir, 1860:279

Distribution: Native, New World. BOLIVIA; BRAZIL; PERÚ. Way of Life: MACROHABITAT: Lowlands to midlands, 204 – 600 meters altitude, in rainforest. MICROHABITAT: Suspended dry leaves in bamboo canopies (Guadua weberbaueri Pilger) at 4m above the ground and in the mid-strata of the rainforest. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in October – November. BEHAVIOR: See description under genus, above. An adult from Perú was collected in a malaise trap, however the specimen is brachypterous. Adults were found in the microhabitat described above using insecticidal fogging techniques. References: Erwin (2001), Erwin (field notes), Naviaux (1998), Pearson, et al. (1999b), Data from D. Pearson, NMNH collection Ctenostoma (Myrmecilla) oblitum Chaudoir 1860 EFFACED COMB-MOUTHED BEETLE Ctenostoma oblitum Chaudoir, 1860:50

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

SPECIES ACCOUNTS / CARABIDAE

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195

Ctenostoma (Naviauxiana) onorei colombiense Naviaux 1998 COLOMBIAN COMB-MOUTHED BEETLE Ctenostoma onorei colombiensis Naviaux, 1998:36

Distribution: Native, New World. COLOMBIA. Way of Life: MACROHABITAT: Uplands to mountains, 2000+ meters altitude, in cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) onorei onorei Naviaux 1998 ONORE’S COMB-MOUTHED BEETLE Ctenostoma onorei onorei Naviaux, 1998:35

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in coastal rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in April. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) ornatum ornatum Klug 1834 ORNATE COMB-MOUTHED BEETLE Ctenostoma ornatum Klug, 1834:42

Distribution: Native, New World. BRAZIL. WAY OF LIFE: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) ornatum werneri Navaiux 1998 WERNER’S COMB-MOUTHED BEETLE Ctenostoma ornatum werneri Navaiux, 1998:51

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica), 60 – 450 meters altitude. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January – February, and November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

196

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Ctenostoma (Naviauxiana) panamense Naviaux & Brzoska 2005 PANAMANIAN COMB-MOUTHED BEETLE Ctenostoma panamensis Naviaux & Brzoska, 2005:21

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Midlands, 550 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: The adult holotype was found in June. BEHAVIOR: See description under genus, above. References: Naviaux & Brzoska (2005) Ctenostoma (Paractenosoma) parallelum Navaiux 1998 PARALLEL-SIDED COMB-MOUTHED BEETLE Ctenostoma parallelum Navaiux, 1998:172

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES : Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) parvulum Navaiux 1998 DIM COMB-MOUTHED BEETLE Ctenostoma parvulum Navaiux, 1998:157

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in November and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) pearsoni Navaiux 1998 PEARSON’S COMB-MOUTHED BEETLE Ctenostoma pearsoni Navaiux, 1998:57

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Lowlands, 50 – 100 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, capable of flight. SEASONAL OCCURRENCE: Adults have been found in April – July. BEHAVIOR: See description under genus, above. Adults are attracted to lights at night. References: Erwin (2001), Naviaux (1998)

SPECIES ACCOUNTS / CARABIDAE

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197

Ctenostoma (Myrmecilla) plicaticolle W. Horn 1911 PLEATED-COLLAR COMB-MOUTHED BEETLE Ctenostoma plicaticolle W. Horn, 1911:153

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Lowlands, 450 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in October – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Microprocephalus) pusillum minimum Navaiux 1998 TINY-TINY COMB-MOUTHED BEETLE Ctenostoma pusillum minimum Navaiux, 1998:86

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Lowlands, 220 meters altitude, in rainforest. MICROHABITAT: The adult holotype was collected from the rainforest canopy using insecticidal fogging techniques. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in October. BEHAVIOR: See description under genus, above. Adults have been commonly found using insecticidal fogging techniques. References: Erwin (2001), Erwin et al. (2004), Naviaux (1998), Pearson et al. (1999a), Data from NMNH collection Ctenostoma (Microprocephalus) pusillum pusillum Navaiux 1998 TINY COMB-MOUTHED BEETLE Ctenostoma pusillum pusillum Navaiux, 1998:85

Distribution: Native, New World. FRENCH GUIANA. Way of Life: MACROHABITAT: Lowlands, 50 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May, July – August, and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) pygmaeum concolor Naviaux 1998 UNIFORM SMALL COMB-MOUTHED BEETLE Ctenostoma pygmaeum concolor Naviaux, 1998:155

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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Ctenostoma (Myrmecilla) pygmaeum pygmaeum (Lacordaire) 1843 PIGMY COMB-MOUTHED BEETLE Myrmecilla pygmaea Lacordaire, 1843:120

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) rapillyi Naviaux 1998 RAPILLY’S COMB-MOUTHED BEETLE Ctenostoma rapillyi Naviaux, 1998:112

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) regium Naviaux 1998 REGAL COMB-MOUTHED BEETLE Ctenostoma regium Naviaux, 1998:123

Distribution: Native, New World. ECUADOR; PERÚ. Way of Life: MACROHABITAT: Lowlands, 200 – 300 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in November – December. BEHAVIOR: See description under genus, above. According to Pearson, et al. (1999a), “Unlike almost all other species of this genus in Ecuador, the eyes are not distinctively wider than the rest of the head. The pronotum is distinctly rounded and domed. The apex of each elytron has a shallow scallop-shaped notch. The majority of these peculiar characters are apparently to enhance mimicry of the powerfully stinging ant Paraponera [clavata (Fabricius) 1775)]. Not only is this beetle the same size and shape as this large ant, but it walks and behaves similarly, even to the point of producing stridulation sounds of the same wave length frequency as the ant (P. DeVries, pers. comm.). This species of tiger beetle is known from the understory vegetation of primary forest in the lowlands of eastern Ecuador and Peru.” References: Erwin (2001), Naviaux (1998), Pearson et al. (1999a), Data from D. Pearson collection

SPECIES ACCOUNTS / CARABIDAE

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199

Ctenostoma (Myrmecilla) rivalieri Naviaux 1998 RIVALIER’S COMB-MOUTHED BEETLE Ctenostoma rivalieri Naviaux, 1998:132

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica), 2500 meters altitude. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April – June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) rugicolle W. Horn 1904 ROUGH-NECK COMB-MOUTHED BEETLE Ctenostoma Batesi rugicollis W. Horn, 1904:82

Distribution: Native, New World. BRAZIL; ECUADOR; PERÚ. Way of Life: MACROHABITAT: Lowlands to mountains, 400 – 2228 meters altitude, in rainforest, cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January and September – October. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Pearson et al. (1999a) Ctenostoma (Ctenostoma) rugiferum pseudoheydeni W. Horn 1913 FALSE HEYDEN’S COMB-MOUTHED BEETLE Ctenostoma rugifrons pseudoheydeni W. Horn, 1913:2

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) rugiferum rugiferum (W. Horn) 1895 WRINKLED COMB-MOUTHED BEETLE Procephalus rugifer W. Horn, 1895:354

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

200

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A TREATISE ON THE WESTERN HEMISPHERE CARABOIDEA. VOLUME I

Ctenostoma (Euctenostoma) rugosum Klug 1824 FOLDED COMB-MOUTHED BEETLE Ctenostoma rugosum Klug, 1824:7 Ctenostoma Dimidiatum Dejean, 1837:6

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, wing length unfolded about that of elytron, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) sahlbergi Chaudoir 1860 SAHLBERG’S COMB-MOUTHED BEETLE Ctenostoma Sahlbergi Chaudoir, 1860:281

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) sallei Chaudoir 1860 SALLE’S COMB-MOUTHED BEETLE Ctenostoma Sallei Chaudoir, 1860:275

Distribution: Native, New World. VENEZUELA. Way of Life: MACROHABITAT: Unknown. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) schaumi pilosulum W. Horn 1898 FELT-COVERED COMB-MOUTHED BEETLE Ctenostoma pilosulum W. Horn, 1898:29

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Alantic forest (Mata Atlântica). The type locality of “pilosulum” was Mata Atlântica when the type specimen was collected. At present, that part of the tropical moist forest in Pernambuco is mostly cultivated sugar cane fields. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in January – March. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

SPECIES ACCOUNTS / CARABIDAE

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201

Ctenostoma (Myrmecilla) schaumi schaumi W. Horn 1895 SCHAUM’S COMB-MOUTHED BEETLE Ctenostoma Schaumi W. Horn, 1895:354

Distribution: Native, New World. BRAZIL, FRENCH GUIANA. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005) Ctenostoma (Myrmecilla) schmalzi paraguayense Brouerius van Nidek 1956 PARAGUAYAN COMB-MOUTHED BEETLE Ctenostoma paraguayense Brouerius van Nidek, 1956:320

Distribution: Native, New World. PARAGUAY. Way of Life: MACROHABITAT: South American Campos ecosystem, gallery forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) schmalzi schmalzi W. Horn 1898 SCHMALZ’S COMB-MOUTHED BEETLE Ctenostoma Schmalzi W. Horn, 1898:32

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) simile Naviaux 1998 VERY SIMILAR COMB-MOUTHED BEETLE Ctenostoma simile Naviaux, 1998:43

Distribution: Native, New World. COSTA RICA; PANAMÁ. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 850 meters, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March and June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection

202

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Ctenostoma (Paractenosoma) simpliceps W. Horn 1900 SIMPLE-HEADED COMB-MOUTHED BEETLE Ctenostoma simpliceps W. Horn, 1900:361

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) spinosum Naviaux 1998 SPINY COMB-MOUTHED BEETLE Ctenostoma spinosum Naviaux, 1998:49

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Midlands, 600 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Naviauxiana) subtilesculptum W. Horn 1913 SUBTLY-SCULPTED COMB-MOUTHED BEETLE Ctenostoma ebenium subtilesculpta W. Horn, 1913:1

Distribution: Native, New World. BOLIVIA; BRAZIL. Way of Life: MACROHABITAT: Lowlands to midlands, 410 – 750 meters altitude, tropical dry forest, cerrados in Brazil. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) succinctum (Laporte de Castelnau) 1834 VERY SHORT COMB-MOUTHED BEETLE Procephalus succinctus Laporte de Castelnau, 1834:36

Distribution: Native, New World. ECUADOR, FRENCH GUIANA; PERÚ, VENEZUELA. Way of Life: MACROHABITAT: Lowlands to mountains, 290 – 2228 meters altitude, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in March and November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Pearson et al. (1999a)

SPECIES ACCOUNTS / CARABIDAE

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203

Ctenostoma (Euctenostoma) sumlini Naviaux 1998 SUMLIN’S COMB-MOUTHED BEETLE Ctenostoma sumlini Naviaux, 1998:118

Distribution: Native, New World. PERÚ. Way of Life: MACROHABITAT: Midlands, 664 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in September. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Ctenostoma) transversum Naviaux 1998 TRANSVERSELY-MARKED COMB-MOUTHED BEETLE Ctenostoma transversum Naviaux, 1998:103

Distribution: Native, New World. BOLIVIA; BRAZIL. Way of Life: MACROHABITAT: Lowlands, 50 – 310 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January, and October – December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) trinotatum (Fischer von Waldheim) 1821 THREE-SPOTTED COMB-MOUTHED BEETLE Caris fasciatum Fischer von Waldheim, 1821: pl. 1 Caris trinotata Fischer von Waldheim, 1821:99 Ctenostoma Fischeri Chaudoir, 1848:22

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) tumidum Naviaux 1998 SWOLLEN-COLLARED COMB-MOUTHED BEETLE Ctenostoma tumidum Naviaux, 1998:75

Distribution: Native, New World. COSTA RICA. Way of Life: MACROHABITAT: Midlands, 1280 meters altitude, in cloudforests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in June. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Data from NMNH collection

204

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Ctenostoma (Neoprocephalus) turnbowi Naviaux 1998 TURNBOW’S COMB-MOUTHED BEETLE Ctenostoma turnbowi Naviaux, 1998:82

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Midlands to uplands, 1000 – 2000 meters altitude, in cloudforests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Euctenostoma) tyrannum (Thomson) 1859 MASTER COMB-MOUTHED BEETLE Procephalus tyrannus Thomson, 1859:92

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) unifasciatum affine W. Horn 1894 KIN COMB-MOUTHED BEETLE Ctenostoma affine W. Horn, 1894:109 Ctenostoma Gautardi affine W. Horn, 1908:91

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) unifasciatum unifasciatum Dejean 1831 ONE-LINED COMB-MOUTHED BEETLE Ctenostoma trinotatum Klug, 1824:5 Ctenostoma Unifasciatum Dejean, 1831:272 Ctenostoma trifasciatum Lacordaire, 1843:118 Ctenostoma Klugii Lacordaire, 1843:119 Ctenostoma laceratum R.F. Sahlberg, 1844:503

Distribution: Native, New World. BRAZIL.

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Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in April and November. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) vairai Cassola 2001 VAIRA’S COMB-MOUTHED BEETLE Ctenostoma vairai Cassola, 2001:61

Distribution: Native, New World. ECUADOR. Way of Life: MACROHABITAT: Midlands, 1400 meters altitude, in lower montane rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in February. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) vicinum Naviaux 1998 NEIGHBORHOOD COMB-MOUTHED BEETLE Ctenostoma vicinum Naviaux, 1998:136

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Neoprocephalus) wappesi Naviaux 1998 WAPPES’ COMB-MOUTHED BEETLE Ctenostoma wappesi Naviaux, 1998:79

Distribution: Native, New World. PANAMÁ. Way of Life: MACROHABITAT: Lowlands to midlands, 50 – 850 meters altitude, in rainforest and cloud forests. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in May – July. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998), Naviaux & Brzoska (2005), Data from CAS collection

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Ctenostoma (Euctenostoma) wiesneri Naviaux 1998 WIESNER’S COMB-MOUTHED BEETLE Ctenostoma wiesneri Naviaux, 1998:114

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Adults have been found in December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) zerchei Naviaux 1998 ZERCHE’S COMB-MOUTHED BEETLE Ctenostoma zerchei Naviaux, 1998:131

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica). MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Salvioides) zikani W. Horn 1911 ZIKAN’S COMB-MOUTHED BEETLE Ctenostoma Zikani W. Horn, 1911:152

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Southern Atlantic Forest (Mata Atlântica), 611 meters altitude. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Macropterous, probably capable of flight. SEASONAL OCCURRENCE: Adults have been found in January and December. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998) Ctenostoma (Myrmecilla) zonata Chaudoir 1860 GIRDLED COMB-MOUTHED BEETLE Ctenostoma zonata Chaudoir, 1860:279

Distribution: Native, New World. BRAZIL. Way of Life: MACROHABITAT: Lowlands, 200 meters altitude, in rainforest. MICROHABITAT: Arboreal, on twigs and leaves above the ground. DISPERSAL ABILITIES: Brachypterous, hence flightless thus vagility limited to walking or running. SEASONAL OCCURRENCE: Unknown. BEHAVIOR: See description under genus, above. References: Erwin (2001), Naviaux (1998)

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General Bibliography for Caraboidea of the Western Hemisphere ACCIAVATTI, R.E. (1980) A review of Cicindela praetextata from the southwest United States (Coleoptera: Cicindelidae). Southwestern Entomologist, 5(4), 231-244. ACLOQUE, A. (1896) Faune de France, contennant la description des espéces indigènes disposées en tableaux analyiques et illustrées de figures représentant les types caractéristiques des genres. 1. Coléoptères. 466pp. J.B. Ballière & Fils, Paris. ÁDÁM, L. (1996) A check-list of the Hungarian caraboid beetles (Coleoptera). Folia Entomologica Hungarica, Rovartani Közlemények (S.N.) 57, 5-64. ADAMS, M.F. (1812) Description de trois coléoptères inconnus de la Sibérie orientale. Mémoires de la Société Impériale des Naturalistes de Moscou, 3, 165-172. ADAMS, M.F. (1817) Descriptio insectorum novorum Imperii Russici, inprimus Caucasi et Sibiriae. Mémoires de la Société Impériale des Naturalistes de Moscou, 5, 278- 314. ADIS, J. (1981) Systematics and natural history of Solenogenys Westwood (Coleoptera: Carabidae: Scaritini) with a description of a new species from the central Amazon, Brazil. The Coleopterists Bulletin, 35(2), 153-166. AGASSIZ, J.L.R. (1846) Nomenclator zoologicus. Fasciculus XI. Continens Coloeptera. 170pp. Soloduri, Switzerland. AGASSIZ, J.L.R. (1847) Nomenclatoisr Zoologici. Index Universalis …393pp. Jenet et Gassmann, Soloduri, Switzerland. AHRENS, A. (1810) Beiträge zu einer Monographie der deutschen Rohrkäfer-Arten. Neueste Schriften Naturforschenden Gesellschaft zu Halle, 1(3), 1-48. AHRENS, A. (1812) Beitrag zur Kenntniss deutscher Käfer. Neue Schriften Naturforschenden Gesellschaft zu Halle, 2(2), 1-40. AHRENS, A. (1830) Beschreibung einiger deutscher Arten der Gattung Clivina. Thon’s Entomologisches Archiv, 2, 57-61. ALCOCK, J. (1976) The behaviour of the seed-collecting larvae of a carabid beetle (Coleoptera). Journal of Natural History 10, 367-375. ALEXANDER, RD. 1959. The courtship and copulation of Pasimachus punctulatus Haldeman (Coleoptera: Carabidae). Annals of the Entomological Society of America, 52, 485. ALLEN, R.T. (1968) A synopsis of the Tribe Morionini in the Western Hemisphere with descriptions of two new species (Coleoptera: Carabidae). Caribbean Journal of Science, 8(3-4), 141-163. ALLEN, R.T. (1972) A revision of the Genus Loxandrus LeConte (Coleoptera: Carabidae) in North America. Entomologica Americana, 46(1), 1-184. ALLEN, R.T. (1973) Peronoscelis Chaudoir known from Bolivia, with a description of a new species (Coleoptera: Carabidae). Revista Brasileira de Entomologia, 17(9), 57-63. ALLEN, R.T. (1977) Calosoma (Castrida) alternans granulatum Perty: A predator of cotton leaf worms in Bolivia (Coleoptera: Carabidae: Carabini). The Coleopterists Bulletin, 31(1), 73-76. ALLEN, RT. (1979) The occurrence and importance of ground beetles in agricultural and surrounding habitats. pp. 485-505. In, Erwin, T.L., Ball, G.E., Whitehead, D.R. & Halpern,

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A.L. (eds.), Carabid Beetles: Their Evolution, Natural History, and Classification, Proceedings of the First International Symposium of Carabidology. Dr. W. Junk, Publishers, The Hague. ALLEN, R.T. (1980) A review of the subtribe Myadi: description of a new genus and species, phylogenetic relationships, and biogeography (Coleoptera: Carabidae: Pterostichini). The Coleopterists Bulletin, 34, 1-29. ALLEN, R.T. & BALL, G.E. (1980) Synopsis of Mexican taxa of the Loxandrus series (Coleoptera: Carabidae: Pterostichini). Transactions of the American Entomological Society, 105, 481-576. ALLEN, R.T. & CARLTON, L.E. (1988) Two new Scaphinotus from Arkansas with notes on other Arkansas species (Coleoptera: Carabidae: Cychrini). Journal of the New York Entomological Society, 96(2), 129-139. ALLEN, R T. & THOMPSON, R.G. (1977) Faunal composition and seasonal activity of Carabidae (Insecta: Coleoptera) in three different woodland communities in Arkansas. Annals of the Entomological Society of America, 70, 31-34. ALLUAUD, C. (1919) Contributions à l’étude des Carabiques d’Afrique et de Madagascar [Col.] XXIII. Observations sur divers clivindes; descriptions d’une race, d’une espèce et d’un genre nouveaux. Bulletin de la Société Entomologique de France, 1919, 99-102. ALLUAUD, C. (1926) Note sur les carabiques. III. Description d’un Bembidion des Açores et notes sur la faune des îles de l’Atlantique. Bulletin de la Société Entomologique de France, 1926, 11-13. ALLUAUD, C. (1932) Carabiques du Maroc. Afra, Cahiers d’Entomologie, 5, 1-4. ALVARES, E.S.S. & FERREIRA, R.L. (2002) Coarazuphium pains, a new species of troglobitic beetle from Brazil (Coleoptera: Carabidae: Zuphiini). Lundiana, 3(1), 41-43. ANDERSEN, J. (1968) The larva of Miscodera arctica Payk. (Co1. Carabidae). Norsk Entomologisk Tidsskrift, 15, 71-74. ANDREWS, H.E. (1921) Notes on Oriental Carabidae, II. Some new species of Bembidium from Java and Indo-china. The Entomologist’s Monthly Magazine, 57, 248-252. ANDREWS, H.E. (1924) Papers on Oriental Carabidae. — XIII. Annals and Magazine of Natural History (9) 14, 241-244. ANDREWS, H.E. (1931) Coleoptera-Carabidae of the Juan Fernandez Islands, pp. 629-637. In, Skottsberg, C. (Ser. Ed.), Natural History of the Juan Fernandez and Easter Island, Vol. 3, Uppsala, Aluquist Wiksells Boktyckeri. ANDREWS, H.E. (1937) On some new species of Carabidae, chiefly from Java. (III). Treubia, 16, 25-35. ANTOINE, M. (1943) Notes d’ entomologie marocaine. XL. Sur quelques Carabiques marocaine. Bulletin de la Société d’Histoire Naturelle d’ Maroc, 23, 43-55. ANTOINE, M. (1949) Notes d’entomologie marocaine. L. Sur les Carabiques recueillis au Maroc par MM. Paulian et Villiers. Revue Française d’Entomologie, 16(1), 65-69. ANTOINE, M. (1951) Notes d’entomologie marocaine. LII. Coléoptères nouveaux ou intéressants. Bulletin de la Société d’Histoire Naturelle d’ Maroc, 29(1949), 311-321. ANTOINE, M. (1956) Coléoptères carabiques du Maroc, première partie. Mémoires de la Société des Sciences naturelle et Physiques du Maroc, Rabat, N.S. (Zool.) 1(1955), 1-177. ANTOINE, M. (1961) Coléoptères carabiques du Maroc. Quatrième partie. Mémoires de la Société des Sciences naturelle et Physiques du Maroc, Rabat, N.S. (Zool.) 8, 467-537. APFELBECK, V. (1904) Die Käferfauna der Balkanhalbinsel, mit Berücksichtigung Klein-Asiens und der Insel Kreta. Erster Band. Familienreihe Caraboidea. ix + 422pp. R. Friedländer und Sohn, Berlin. APFELBECK, V. (1918) Koleopteren aus dem nordalbanish-montenegrinischen Grenzgebiete. Ergebnisse einer von der Kaiserl. Akademie der Wissenschaften in Wien veranlassten naturwissenschftlichen Forschungsreise in Nordabanien [sic]. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-naturwissenschaftliche Klasse, 127, 159-176.

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ARNDT, E. (1998) The species of Notiobia Perty (Coleoptera: Carabidae: Harpalini) from Brazil. Acta Amazonica, 28(3), 285-299. ARNDT, E. & WRASE, D.W. (2001) Description of two new species of Notiobia Perty from southern Venezuela. (Insecta, Coleoptera, Carabidae, Harpalini). Spixiana, 24(2), 157-163. ARROW, G.J. (1942) The beetle family Rhysodidae, with some new species and a key to those at present known. Proceedings of the Royal Entomological Society of London, (B), 11, 171-183. AUDINET-SERVILLE, J.G. (1821) Fauna Française ou histoire naturelle, générale et particulière des animaux qui se trouvent en France, constamment ou passagèrement, à la surface du sol, dans les eaux qui le baignent, et dans le littoral des mers qui le bornent, Coléoptères. 240pp. Levault, Paris. AUDUOIN, J.V. (1836) IN, DEJEAN, P.F.M.A. (1836) Catalogue des coléoptères de la collection de M. le Comte Dejean, Troisième edition, revue, corrigée et augmentée, Paris. 468 pp. AUDUOIN, J.V. & BRULLÉ, G.A. (1839) Description des espèces nouvelles ou peu connues de la famille de cicindelètes faisant partie de la collection du Muséum. Archives du Muséum d’Histoire Naturelle, 1, 115-142. AVON, C. (1994) Un nouveau Peryphus de France localisé dans les Alpes-Maritimes (Coleoptera, Carabidae, Bembidiitae, Bembidiini). L’Entomologiste (Paris), 50(3), 157-160. BAEHR, M. (1983a) Pericompsus striatopunctatus sp. nov. aus dem brasilianischen Amazonasgebiet. Spixiana, 6(1), 83-85. BAEHR, M. (1983b) Schizogenius freyi sp. nov., die erste Schizogenius-Art ausserhalb Amerikas (Coleoptera, Carabidae, Scarabinae). Entomologische Arbeiten aus dem Museum Georg Frey, 3132, 1983, 91-95. BAEHR, M. (1997a) Two new Pseudomigadops Jeannel, 1938 from the Falkland Islands (Coleoptera, Carabidae, Migadopinae). Mitteilungen der Münchener Entomologischen Gesellchaft, 87, 39-45. BAEHR, M. (1997b) Revision of the Pseudomorphinae of the Australian region. 2. The genera Pseudomorpha Kirby, Adelotopus Hope, Cainogenion Notman, Paussotropus Waterhouse, and Cryptocephalomorpha Ritsema. Taxonomy, phylogeny, zoogeography. (Insecta, Coleoptera, Carabidae). Spixiana, Supplement 23, 1-508. BAEHR, M. (1999) Further notes on Migadopinae from the Falkland Islands (Insecta, Coleoptera, Carabidae). Spixiana, 22(1), 47-52. BAEHR, M. (2001) Syntopic and sycnchronic occurrence of closely related species of the genus Scarites Fabricius in Amazonian Brazil (Insecta, Coleoptera, Carabidae, Scaritinae). Spixiana, 25(3), 225-237. BALIANI, A. (1934) Studi sulle Amara asiatiche (Col., Carab.) con descrizione di nuove specie. Memorie della Società Entomologica Italiana, 12 (1933), 188-208. BALIANI, A. (1936) Studi sulle Amara asiatiche (Coleopt., Carab.) X. Memorie della Società Entomologica Italiana, 15, 176-183. BALIANI, A. (1938) Studi sulle Amara asiatiche (Coleopt., Carab.) XI. Bollettino della Società Entomologica Italiana, 70, 183-187. BALIANI, A. (1939a) Amara nuove d’Africa e d’Europa. Bollettino della Società Entomologica Italiana, 71(1), 14-16. BALIANI, A. (1939b) Studi sulle Amara asiatiche (Coleopt., Carab.) XII. Bollettino della Società Entomologica Italiana, 71, 136-138. BALIANI, A. (1943) Studi sulle Amara asiatiche XIV (Coleopt., Carabidae). Memorie della Società Entomologica Italiana (Genova), 22(1), 38-50. BALL, G.E. (1951) A note concerning the correct application of the generic name Helluomorpha Castelnau. Bulletin of the Brooklyn Entomological Society, 46: 135-136.

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BALL, G.E. (1956) A revision of the North American species of the Genus Helluomorphoides Ball, 1951 (Coleoptera: Carabidae: Helluonini). Proceedings of the Entomological Society of Washington, 58(2), 67-91. BALL, G.E. (1959) A taxonomic study of the North American Licinini with notes on the Old World species of the Genus Diplocheila Brullé (Coleoptera). Memoirs of the American Entomological Society, 16, iv + 1-258. BALL, G. E. 1962. Descriptions of eleven new species of the beetle subgenus Cryobius (Pterostichus, Col. Carabidae) from Alaska and northwestern Canada. Opuscula Entomologica, 28[1963], 1-26. BALL, G.E. (1966a) The taxonomy of the subgenus Scaphinotus Dejean with particular reference to the subspecies of Scaphinotus petersi Roeschke (Coleoptera: Carabidae: Cychrini). Transactions of the American Entomological Society, 92, 687-722. BALL, G.E. (1966b) A revision of the North American species of the subgenus Cryobius Chaudoir (Pterostichus, Carabidae, Coleoptera). Opuscula Entomologica, Supplementum XXVIII, 1-166. BALL, G.E. (1970) The species of the Mexican genus Aztecarpalus new genus (Coleoptera: Carabidae: Harpalini). The Coleopterists Bulletin, 24, 97-123. BALL, G.E. (1972) Classification of the species of the Harpalus subgenus Glanodes Casey (Carabidae, Coleoptera). The Coleopterists Bulletin, 26, 179-204. BALL, G.E. (1973) A new name for Harpalus (Glanodes) punctieps (Casey) (Coleoptera: Carabidae). The Coleopterists Bulletin, 27, 74. BALL, G.E. (1975a) Pericaline Lebiini: Notes on classification, a synopsis of the New World genera, and a revision of the Genus Phloeoxena Chaudoir (Coleoptera: Carabidae). Quaestiones entomolgicae, 11, 143-242. BALL, G.E. (1975b) Phloeoxena newtoni, new species, and notes on P. nigricollis Ball and P. geniculata Chaudoir, from Mexico. Entomological News, 86, 151-156. BALL, G.E. (1976) Aztecarpalus Ball: new species from Oaxaca, Mexico, reclassification, and a reconstructed phylogeny of the hebescens Group (Coleoptera: Carabidae: Harpalini). The Coleopterists Bulletin, 30, 61-72. BALL, G.E. (1978) The species of the Neotropical Genus Trichopselaphus Chaudoir (Coleoptera: Carabidae: Harpalini): Classification, phylogeny and zoogeography. Quaestiones entomolgicae, 14, 447-489. BALL, G.E. (1987) The Neotropical Genus Trichopselaphus Chaudoir (Coleoptera: Carabidae: Harpalini): New species and a review of taxonomic and evolutionary aspects. Transactions of the American Entomological Society, 112, 249-287. BALL, G.E. (1991) Taxonomic notes about the Middle American genus Cyrtolaus Bates (Coleoptera: Carabidae: Pterostichini). Proceedings of the Entomological Society of Washington, 93, 230-239 BALL, G.E. (1992) The tribe Licinini (Coleoptera: Carabidae): a review of the genus- groups and of the species of selected genera. Journal of the New York Entomological Society, 100, 325-380. BALL, G.E. (2000) Notes about the Genus Tetragonoderus Dejean with description of T. (Peronoscelis) matilei a new species from the Amazon Basin (Coleoptera: Carabidae: Cyclosomini). Revue Française d’Entomologie, [N.S.] 22(4), 189196. BALL, G.E. (2001) The Subgenera of Clivina Latreille in the Western Hemisphere, and a revision of Subgenus Antroforceps Barr (new status), with notes about evolutionary aspects (Coleoptera: Carabidae: Clivinini). Special publications of the Japan Coleoptera Society, Osaka, 1, 129-156. BALL, G.E. & ANDERSON, J.N. (1962) The taxonomy and speciation of Pseudophonus. The Catholic University of America Press, Washington, D.C. 99 pp. BALL, G.E. & ERWIN, T.L. (1969) A synopsis of the Tribe Loricerini (Coleoptera: Carabidae). Canadian Journal of Zoology, 47(5), 877907.

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BALL, G.E. & HILCHIE, G. J. (1983) Cymindine Lebiini of authors: redefinition and reclassification of genera (Coleoptera: Carabidae). Quaestiones Entomologicae, 19, 93-216. BALL, G.E. & MADDISON, D.R. (1987) Classification and evolutionary aspects of the species of the New World Genus Amblygnathus Dejean, with description of Platymetopsis, new genus, and notes about selected species of Selenophorus Dejean Chaudoir (Coleoptera: Carabidae: Harpalini). Transactions of the American Entomological Society, 113, 189-307. BALL, G.E. & MCCLEVE, S. (1990) The Middle American genera of the Tribe Ozaenini with notes about the speices in southwestern United States and selected species from Mexico. Quaestiones entomologicae, 26, 30-116. BALL, G.E. & NÈGRE, J. (1972) The taxonomy of the Nearctic species of the genus Calathus Bonelli (Coleoptera: Carabidae: Agonini). Transactions of the American Entomological Society, 98, 412-533. BALL, G.E. & NIMMO, A.P. (1983) Synopsis of the species of Subgenus Progaleritina Jeannel, including reconstructed phylogeny and geographical history (Coleoptera: Carabidae: Galerita Fabricius). Transactions of the American Entomological Society, 109, 295-356. BALL, G.E. & ROUGHLEY, R. (1982) The Hypherpes-like taxa of southern Mexico: classification and evolutionary considerations (Coleoptera: Carabidae: Pterostichus). Transactions of the American Entomological Society, 108: 315-399. BALL, G.E. & SHPELEY D. (1983) The species of eucheiloid Pericalina: Classification and evolutionary considerations (Coleoptera: Carabidae: Lebiini). The Canadian Entomologist, 115, 743-806. BALL, G.E. & SHPELEY D. (1992) Appendix, pp. 94-121. In, Noonan, G.R., Ball, G.E. & Stork, N.E. (eds.), The Biogeography of Ground Beetles of Mountains and Islands. vii + 256 pp. Intercept Ltd., Andover. BALL, G.E. & SHPELEY D. (2002a) Ginemini, Ginema thomasi, New Tribe, New Genus and New Species, from Amazonian Bolivia (Coleoptera: Carabidae: Harpalinae). Transactions of the American Entomological Society, 128(1), 75-98. BALL, G.E. & SHPELEY D. (2002b) The Neotropical Subgenera and species of the Pantropical Genus Anaulacus MacLeay (sensu novo) (Coleoptera: Carabidae: Masoreini): A taxonomic revision, with notes about way of life, evolution, and geographical history. Transactions of the American Entomological Society, 128(2+3), 265-343. BALL, G.E. & SHPELEY D. (2005) Taxonomic review of the Tribe Melaenini (Coleoptera: Carabidae), with observations on morphological, ecological and chorological evolution, Zootaxa 1099, 1-120. BALL, G.E., SHPELEY D. & CURRIE, D.C. (1991) The New World Genus Stenomorphus Dejean (Coleoptera: Carabidae: Harpalini): Classification, allometry, and evolutionary considerations. The Canadian Entomologist, 123, 933-988. BALL, G.E. & BOUSQUET, Y. (2001) Carabidae Latreille, 1810, Chapter 6, pp. 33-132. In, Arnett, R.H. & Thomas, M.C. (eds.) American Beetles, 1. CRC Press, Boca Raton, London, New York, Washington, D.C. BALL G.E. & SHPELEY, D. (2002) A taxonomic review of species of the Neotropical pericaline Genus Oreodicastes Maindron, 1905 (Coleoptera: Carabidae: Lebiini). Insecta Mundi, (2001), 15(2), 71-76. BALLION, E. (1870) Eine Centurie neuer Käfer aus der Fauna der russischen Reiches. Bulletin de la Société Impériale des Naturalistes de Moscou, 43(1), 320-353. BÄNNINGER, M. (1925) Neunter Beitrag zur Kenntnis der Carabinae: die Nebriini. Entomologische Mitteilungen, 14(2), 180-195; 14(3/4), 256-281; 14(5/6), 329-343. BÄNNINGER, M. (1927a) Die Ozaenini (Col. Carab.). 10. Beitrag zur Kenntnis der Carabinae. Deutsche Entomologische Zeitschrift, 1927, 177-216.

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WHITEHEAD, D.R. & BALL, G.E. (1975) Classification of the genus Cyrtolaus Bates (Coleoptera: Carabidae: Pterostichini). Quaestiones Entomologicae, 11, 596-621. WHITEHEAD, D.R. & BALL, G.E. (1997) The Middle American Genus Onypterygia Dejean (Insecta: Coleoptera: Carabidae: Platynini): A taxonomic revision of the species, with notes about their way of life and geographical distribution. Annals of Carnegie Museum, 66(3), 289-409. WHITEHEAD, D.R. & REICHARDT, H. (1977) Classification of Listropus Putzeys, a subgenus of Schizogenius Putzeys (Coleoptera: Carabidae: Scaritini). The Coleopterists Bulletin, 31(3), 239-250. WICKHAM, H.F. (1897) The Coleoptera of the Lower Rio Grande Valley. Bulletin from the Laboratories of Natural History of the State University of Iowa, 4(2), 109-110. WICKHAM, H.F. (1899) The habits of American Cicindelidae. Proceedings of the Davenport Academy of Natural Sciences, 7, 206-228. WICKHAM, H.F. (1905) New species of Coleoptera from the western United States. The Canadian Entomologist, 37, 165-171. WICKHAM, H.F. (1919a) Two species of Asaphidion from North America. Proceedings of the Entomological Society of Washington, 21, 178-181. WICKHAM, H.F. (1919b) Scaphinotus (Pseudonomaretus) mannii n. sp. (Coleoptera, Carabidae). Proceedings of the Entomological Society of Washington, 21, 170-173. WIESNER, J. (1999a) The Tiger beetle genus Oxycheila (Insecta: Coleoptera: Cicindelidae). 50th Contribution towards the knowledge of Cicindelidae. Coleoptera, 3, 1-81. WIESNER, J. (1999b) Pentacomia cupricollis and descriptions of two related species from Bolivia (Coleoptera, Cicindelidae). Mitteilungen Muenchener Entomologischen Gesellschaft, 89, 87-94. WILL, K. (1998) A new species of Diplocheila Brulle from North America, with notes on female reproductive tract characters in selected Licinini and implications for evolution of the subgenus Isorembus Jeannel (Coleoptera: Carabidae: Licinini). Proceedings of the Entomological Society of Washington, 100(1), 95-103. WILL, K.W. (2001) A new species of Harpalus Latreille (Coleoptera: Carabidae) from southeastern North America. The Coleopterist Bulletin, 55(4), 447-452. WILL, K.W. (2002) Revision of the new world abariform genera Neotalus N. GEN. and Abaris Dejean (Coleoptera: Carabidae: Pterostichini (Auctorum). Annals of the Carnegie Museum of Natural History, 71, 143-213. WILL, K.W. (2004a) A remarkable new species of Trirammatus Chaudoir (Coleoptera: Carabidae: Pterostichini) from the Valdivian Forest of Chile. Zootaxa, 758, 1-9. WILL, K.W. (2004b) New species of the South American loxandrine genus Metoncidus Bates (Coleoptera, Carabidae). Revista Brasileira de Entomologia, 48(4), 441-446. WILL, K.W. (2004c) Review and cladistic analysis of the generic-level taxa of Morionini Brullé (Coleoptera: Carabidae). The Pan-Pacific Entomologist, 79(3/4), 212-229. WILL, K.W. (2005a) New tribal and generic placement for taxa of Pterostichini (auct.) (Coleoptera: Carabidae) from the Juan Fernández Archipelago, Chile with taxonomic notes on Trirammatus Chaudoir, 1838. The Pan-Pacific Entomologist, 81 (112), 68-75. WILL, K.W. (2005b) The Neotropical genera Oxycrepis Reiche and Stolonis Motschulsky: a taxonomic review, key to the described species and description of new Stolonis species from Ecuador (Coleoptera: Carabidae: Loxandrini). Zootaxa, 1049, 1-17. WILL, K. & LIEBHEER, J.K. (1997) New and little known species of Loxandrus LeConte 1852 (Coleoptera: Carabidae) from North and South America. Studies on Neotropical Fauna and Environment, 32, 230-238. WILLIS, H.L. (1967) Bionomics and zoogeography of tiger beetles of saline habitats in the central United States (Coleoptera: Cicindelidae). The University of Kansas Science Bulletin, 48, 145-313.

GENERAL BIBLIOGRAPHY FOR CARABOIDEA OF THE WESTERN HEMISPHERE

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WIESNER, J. (1981) A new Oxycheila species from Mexiko (Coleoptera: Cicindelinae). Entomologica Basiliensia, 6, 150-153. WIESNER, J. (1999) The Tiger beetle genus Oxycheila (Insecta: Coleoptera: Cicindelidae). 50th Contribution towards the knowledge of Cicindelidae. Coleoptera, 3, 1-81. WIESNER, J. (2003) A new Oxycheila species from Brazil (Coleoptera: Cicindelinae). Entomologische Zeitschrift, 113(6), 166-167. WILL, K.W. (2001) A new species of Harpalus Latreille (Coleoptera: Carabidae) from southeastern North America. The Coleopterist Bulletin, 55(4), 447-452. WILL, K.W. (2002a) Revision of the New World Abariform Genera Neotalus N.GEN. and Abaris Dejean (Coleoptera: Carabidae: Pterostichini (auctorum)). Annals of Carnegie Museum, 71(3), 143-213. WILL, K.W. (2002b) A new tribal placement for Glyptolenus mirabilis (Straneo) (Coleoptera: Carabidae), comb. n., and description of G. straneoi, sp. n. from Ecuador. Studies on Neotropical Fauna and Environment, 37(1), 59-64. WOLCOTT, G.N. (1936) “Insectae Borinquenses.” A revised annaotated check-list of the insects of Porto Rico, with descriptions of some new species. The Journal of the Department of Agriculture of Puerto Rico, 7(1), 1923 (1924), 1-313. WOLLASTON, T.V. (1854) Insecta Maderensia; being an account of the insects of the islands of the Madeiran group. 634pp. London. WOLLASTON, T.V. (1858) Brief diagnostic characters of undescribed Madeiran Insects. Annals and Magazine of Natural History, ser. 3, 1, 18-28, 113-125. WOLLASTON, T.V. (1867) Coleoptera Hesperidum, being an enumeration of the coleopterous insects of the Cape Verde Archipelago. 285pp. London. WRIGHT, J.F. (1939) A new Loxandrus (Coleoptera, Carabidae) from Cincinnati, Ohio. Bulletin of the Brooklyn Entomological Society, 34: 257-258. ZABELLOS, J. (1997) Honduranillus balli: Un Nuevo genero y especie de Anillini (Coleoptera, Caraboidea) de Honduras. Acta Zoológica Mexicana, 71, 33-43. ZABELLOS, J. & MATEU, J. (1997) Dos nuevos Anillinus Casey, 1918 (Coleoptera: Caraboidea: Trechidae) de Brasil. Elytron, 11, 133-143. ZAYAS, F. (1988) Entomofauna Cubana. Orden Coleoptera. 212pp. Habana. ZETTERSTEDT, J.W. (1828) Fauna Insectorum Lapponica, Part I. Xx + 563pp. Libraria Schulziana, Hammone. ZETTERSTEDT, J.W. (1838) Insecta Lapponica, 1(1), 191pp. Leipzig. ZIMMERMANN, C.C.A. (1831) Monographie der Carabiden, 1. Zabroidea, 1-76. ZIMMERMANN, C.C.A. (1832) [Über die bisherige Gattung Amara. Ein Beitrag zu einer künftigen Monographis der hierher gehörenden Thiere.] In, Gistel, J.N.F.X., Faunus. 1(1) 5-40. ZIMMERMANN, C.C.A. (1869) Synonymical notes on Coleoptera of the United States, with descriptions of new species, from the MSS of the late Dr. C. Zimmermann (Editied by John L. LeConte, M.D.). Transactions of the American Entomological Society, 2, 243-259. ZIMMERMANN, C.C.A. (1874) In, LeConte, J.L., Notes on the species of Pasimachus. Bulletin of the Buffalo Society of Natural Sciences. 1, 266-273.

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Index C Callisthenes Fischer von Waldheim 74 Callisthenes (Callistenia) dawsoni Dajoz 76 Callisthenes (Callistenia) dietzi (Schaeffer) 76 Callisthenes (Callistenia) discors (LeConte) 77 Callisthenes (Callistenia) lariversi (Van Dyke) 77 Callisthenes (Callistenia) latipennis (G. Horn) 77 Callisthenes (Callistenia) luxatus (Say) 78 Callisthenes (Callistenia) moniliatus LeConte 79 Callisthenes (Callistenia) monticola (Casey) 79 Callisthenes (Callistenia) oregonus Gidaspow 80 Callisthenes (Callistenia) placerus Gidaspow 80 Callisthenes (Callistenia) schaefferi (Breuning) 81 Callisthenes (Callistenia) subaeneus (Chaudoir) 82 Callisthenes (Callistenia) subasperatus (Schaeffer) 82 Callisthenes (Callistenia) wilkesii LeConte 83 Callisthenes (Chrysostigma) affinis (Chaudoir) 74 Callisthenes (Chrysostigma) ampliator (Bates) 75 Callisthenes (Chrysostigma) calidus (Fabricius) 75 Callisthenes (Chrysostigma) cancellatus (Eschscholtz) 76 Callisthenes (Chrysostigma) lepidus (LeConte) 78 Callisthenes (Chrysostigma) morrisonii (G. Horn) 79 Callisthenes (Chrysostigma) obsoletus (Say) 80 Callisthenes (Chrysostigma) semilaevis (LeConte) 81 Callisthenes (Chrysostigma) simplex (LeConte) 81 Callisthenes (Chrysostigma) tepidus (LeConte) 82 Calopachys Haury 83 Calopachys blaptoides (Putzeys) 83 Calopachys bohnei (Korell & Frisch) 84 Calopachys omiltemius (Bates) 84 Calopachys viridissimus (Haury) 84 Calosoma Weber 85 Calosoma (Blaptosoma) anthracinum anthracinum Dejean 87 Calosoma (Blaptosoma) anthracinum microgonum Bates 87 Calosoma (Blaptosoma) atrovirens atrovirens Chaudoir 88 Calosoma (Blaptosoma) atrovirens explanaticolle Bates 88 Calosoma (Blaptosoma) bridgesi Chaudoir 89 Calosoma (Blaptosoma) chihuahua Gidaspow 90 Calosoma (Blaptosoma) haydeni haydeni G. Horn 96 Calosoma (Blaptosoma) haydeni punctulicolle Bates 96 Calosoma (Blaptosoma) laeve Dejean 97

Calosoma (Blaptosoma) porosifrons Bates 101 Calosoma (Blaptosoma) viridisulcatum laetulum (Jeannel) 107 Calosoma (Blaptosoma) viridisulcatum viridilucens Beheim & Breuning 107 Calosoma (Blaptosoma) viridisulcatum viridisulcatu Chaudoir 107 Calosoma (Callitropa) externum (Say) 92 Calosoma (Callitropa) macra (LeConte) 98 Calosoma (Callitropa) palmeri G. Horn 99 Calosoma (Callitropa) protracta LeConte 102 Calosoma (Calosoma) aurocinctum Chaudoir 89 Calosoma (Calosoma) frigidum Kirby 92 Calosoma (Calosoma) scrutator (Fabricius) 104 Calosoma (Calosoma) splendidum Dejean 104 Calosoma (Calosoma) sycophanta (Linné) 105 Calosoma (Calosoma) wilcoxi LeConte 107 Calosoma (Carabomimus) altipeta (Jeannel) 86 Calosoma (Carabomimus) asper (Jeannel) 88 Calosoma (Carabomimus) bolivianum Géhin 89 Calosoma (Carabomimus) bulleri Breuning 90 Calosoma (Carabomimus) cicatricosum Chaudoir 90 Calosoma (Carabomimus) costipenne Chaudoir 90 Calosoma (Carabomimus) depressicolle Chaudoir 91 Calosoma (Carabomimus) digueti (Lapouge) 91 Calosoma (Carabomimus) diminutum Bates 91 Calosoma (Carabomimus) flohri Bates 92 Calosoma (Carabomimus) gebieni (Breuning) 94 Calosoma (Carabomimus) laevigatum hoegei Breuning 97 Calosoma (Carabomimus) laevigatum laevigatum Chaudoir 97 Calosoma (Carabomimus) morelianum Bates 99 Calosoma (Carabomimus) orizabae (Jeannel) 99 Calosoma (Carabomimus) politum Chaudoir 101 Calosoma (Carabomimus) striatulum Chevrolat 105 Calosoma (Carabosoma) angulatum Chevrolat 86 Calosoma (Carabosoma) eremicola Fall 91 Calosoma (Carabosoma) glabratum Dejean 94 Calosoma (Carabosoma) marginale Casey 98 Calosoma (Carabosoma) parvicolle Fall 100 Calosoma (Carabosoma) peregrinator Guérin-Méneville 100 Calosoma (Carabosoma) prominens LeConte 102 Calosoma (Carabosoma) sponsum Casey 104

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Calosoma (Carabosoma) striatipenne Chaudoir 105 Calosoma (Castrida) abbreviatum Chaudoir 85 Calosoma (Castrida) alternans (Fabricius) 86 Calosoma (Castrida) argentinensis Csiki 87 Calosoma (Castrida) fulgens Chaudoir 93 Calosoma (Castrida) galapageium Hope 93 Calosoma (Castrida) granatense darwinia Van Dyke 94 Calosoma (Castrida) granatense floreanum (Basilewsky) 94 Calosoma (Castrida) granatense granatense Géhin 95 Calosoma (Castrida) leleuporum (Basilewsky) 97 Calosoma (Castrida) linelli Mutchler 98 Calosoma (Castrida) peruviense Mandl 101 Calosoma (Castrida) retusum (Fabricius) 102 Calosoma (Castrida) rufipenne Dejean 103 Calosoma (Castrida) sayi Dejean 103 Calosoma (Castrida) trapezipenne Chaudoir 106 Calosoma (Castrida) vagans Dejean 106 Carabina Latreille 73 Carabini Latreille 73 Carabus Linné 108 Carabus (Archicarabus) nemoralis nemoralis Müller 114 Carabus (Aulonocarabus) truncaticollis truncaticol Eschscholtz 116 Carabus (Carabus) goryi Dejean 111 Carabus (Carabus) granulatus granulatus Linné 111 Carabus (Carabus) granulatus hibernicus Lindroth 112 Carabus (Carabus) vinctus (Weber) 117 Carabus (Diocarabus) chaimssonis Fischer von Waldheim 109 Carabus (Hemicarabus) serratus Say 114 Carabus (Homoeocarabus) maeander maeander Fischer von Waldheim 113 Carabus (Megodontus) vietinghoffi vietinghoffi Adams 117 Carabus (Oreocarabus) taedatus agassii LeConte 115 Carabus (Oreocarabus) taedatus taedatus Fabricius 116 Carabus (Tachypus) auratus Linné 108 Carabus (Tachypus) cancellatus Illiger 109 Carabus (Tanaocarabus) finitimus Haldeman 110 Carabus (Tanaocarabus) forreri forreri Bates 110 Carabus (Tanaocarabus) forreri willi Deuve 110 Carabus (Tanaocarabus) hendrichsi Bolívar y Pieltain 113 Carabus (Tanaocarabus) sylvosus Say 115 Ceroglossina Lapouge 118 Ceroglossus Solier 118 Ceroglossus buqueti andestus Kraatz-Koschlau 118 Ceroglossus buqueti argentinensis Jiroux 119 Ceroglossus buqueti arieli Jiroux 119

Ceroglossus buqueti arriagadai Jiroux 119 Ceroglossus buqueti breuningi Heinz & Jiroux 120 Ceroglossus buqueti buqueti (Laporte de Castelnau) 120 Ceroglossus buqueti chiloensis (Hope) 120 Ceroglossus buqueti cupreus (Faz) 121 Ceroglossus buqueti deuvei Jiroux 121 Ceroglossus buqueti lepidus Kraatz-Koschlau 121 Ceroglossus buqueti lorenzi Jiroux 122 Ceroglossus buqueti magdalenaensis Jiroux 122 Ceroglossus buqueti peladosus Kraatz-Koschlau 122 Ceroglossus buqueti subnitens Kraatz-Koschlau 123 Ceroglossus buqueti sybarita Gerstaecker 123 Ceroglossus chilensis angolicus Kraatz-Koschlau 123 Ceroglossus chilensis chilensis (Eschscholtz) 124 Ceroglossus chilensis colchaguensis (Reed) 124 Ceroglossus chilensis cyanicollis Kraatz 124 Ceroglossus chilensis evenoui Jiroux 125 Ceroglossus chilensis fallaciosus Kraatz 125 Ceroglossus chilensis ficheti Jiroux 125 Ceroglossus chilensis germaini Jiroux 126 Ceroglossus chilensis gloriosus Gerstaecker 126 Ceroglossus chilensis kraatzianus Morawitz 126 Ceroglossus chilensis latemarginatus Kraatz-Koschlau 127 Ceroglossus chilensis legrandi Heinz & Jiroux 127 Ceroglossus chilensis meridionalis Heinz & Jiroux 127 Ceroglossus chilensis mochae (Reed) 128 Ceroglossus chilensis nigritulus Mandl 128 Ceroglossus chilensis pseudopatagoniensis Heinz & Jiroux 128 Ceroglossus chilensis seladonicus Kraatz-Koschlau 129 Ceroglossus chilensis solieri Roeschke 129 Ceroglossus chilensis villaricensis Kraatz-Koschlau 129 Ceroglossus darwini darwini (Hope) 130 Ceroglossus darwini jirouxi Deuve 130 Ceroglossus darwini mequignoni Heinz & Jiroux 130 Ceroglossus darwini patagoniensis Breuning 131 Ceroglossus darwini reedi Morawitz 131 Ceroglossus guerini Géhin 131 Ceroglossus magellanicus araucanus Kraatz-Koschlau 131 Ceroglossus magellanicus boeufi Jiroux 132 Ceroglossus magellanicus bouchardi Jiroux 132 Ceroglossus magellanicus caburgansis Dueve 132 Ceroglossus magellanicus candens Géhin 133 Ceroglossus magellanicus davidsoni Jiroux 133 Ceroglossus magellanicus dolhemi Jiroux 133 Ceroglossus magellanicus gerstaeckeri Morawitz 133 Ceroglossus magellanicus magellanicus Géhin 134 Ceroglossus magellanicus newtoni Jiroux 134

INDEX

Ceroglossus magellanicus ortrudheinzae Heinz & Jirzoux 134 Ceroglossus magellanicus similis Kraatz-Koschlau 135 Ceroglossus ochseni (Germain) 135 Ceroglossus speciosus Gerstaecker 135 Ceroglossus suturalis ancudanus Morawitz 136 Ceroglossus suturalis olivaceus Kraatz-Koschlau 136 Ceroglossus suturalis puxel Jiroux 136 Ceroglossus suturalis suturalis (Fabricius) 137 Cicindini Csiki 61 Cicindis Bruch 62 Cicindis horni Bruch 62 Collyridini Brullé 171 Ctenostoma Klug 172 Ctenostoma (Ctenostoma) acciavatti Naviaux 173 Ctenostoma (Ctenostoma) bahiaense Naviaux 175 Ctenostoma (Ctenostoma) batesii Chaudoir 175 Ctenostoma (Ctenostoma) chaudoiri (W. Horn) 178 Ctenostoma (Ctenostoma) formicarium (Fabricius) 183 Ctenostoma (Ctenostoma) guyanense W. Horn 185 Ctenostoma (Ctenostoma) heydeni W. Horn 185 Ctenostoma (Ctenostoma) jekelii jekelii Chevrolat 188 Ctenostoma (Ctenostoma) jekelii raresculptum W. Horn 189 Ctenostoma (Ctenostoma) rugicolle W. Horn 199 Ctenostoma (Ctenostoma) rugiferum pseudoheydeni W. Horn 199 Ctenostoma (Ctenostoma) rugiferum rugiferum (W. Horn) 199 Ctenostoma (Ctenostoma) succinctum (Laporte de Castlnau) 202 Ctenostoma (Ctenostoma) transversum Naviaux 203 Ctenostoma (Euctenostoma) bifasciatum bifasciatum Dejean 176 Ctenostoma (Euctenostoma) bifasciatum horni Naviaux 176 Ctenostoma (Euctenostoma) bondari W. Horn 176 Ctenostoma (Euctenostoma) eburatum Bates 182 Ctenostoma (Euctenostoma) ferum Naviaux 183 Ctenostoma (Euctenostoma) fryi Chaudoir 184 Ctenostoma (Euctenostoma) inca Naviaux 187 Ctenostoma (Euctenostoma) klugeanum W. Horn 189 Ctenostoma (Euctenostoma) luctuosum Chaudoir 190 Ctenostoma (Euctenostoma) magnum Naviaux 192 Ctenostoma (Euctenostoma) monnei Naviaux 193 Ctenostoma (Euctenostoma) rapillyi Naviaux 198 Ctenostoma (Euctenostoma) regium Naviaux 198 Ctenostoma (Euctenostoma) rugosum Klug 200 Ctenostoma (Euctenostoma) sahlbergi Chaudoir 200 Ctenostoma (Euctenostoma) sumlini Naviaux 203 Ctenostoma (Euctenostoma) trinotatum (Fischer von Waldheim) 203

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Ctenostoma (Euctenostoma) tyrannum (Thomson) 204 Ctenostoma (Euctenostoma) wiesneri Naviaux 206 Ctenostoma (Microprocephalus) brevilabre W. Horn 177 Ctenostoma (Microprocephalus) pusillum minimum Naviaux 197 Ctenostoma (Microprocephalus) pusillum pusillum Naviaux 197 Ctenostoma (Myrmecilla) abbreviatum Naviaux 172 Ctenostoma (Myrmecilla) agnatum agnatum Chaudoir 173 Ctenostoma (Myrmecilla) agnatum fuscum Naviaux 174 Ctenostoma (Myrmecilla) albofasciatum Chaudoir 174 Ctenostoma (Myrmecilla) asperulum Bates 175 Ctenostoma (Myrmecilla) bicristatum Chaudoir 175 Ctenostoma (Myrmecilla) brevicorne W. Horn 177 Ctenostoma (Myrmecilla) breviusculum von Mannerheim 177 Ctenostoma (Myrmecilla) compactum Naviaux 179 Ctenostoma (Myrmecilla) coracinum Naviaux 179 Ctenostoma (Myrmecilla) crucifrons W. Horn 180 Ctenostoma (Myrmecilla) dentifrons W. Horn 180 Ctenostoma (Myrmecilla) dokhturowi W. Horn 181 Ctenostoma (Myrmecilla) flexuosum Naviaux 183 Ctenostoma (Myrmecilla) gautardi Chaudoir 184 Ctenostoma (Myrmecilla) globifrons W. Horn 184 Ctenostoma (Myrmecilla) hirsutum W. Horn 185 Ctenostoma (Myrmecilla) ichneumoneum ellipticum Naviaux 186 Ctenostoma (Myrmecilla) ichneumoneum ichneumoneum Dejean 187 Ctenostoma (Myrmecilla) infimum Naviaux 187 Ctenostoma (Myrmecilla) luteum Naviaux 191 Ctenostoma (Myrmecilla) macilentum Klug 191 Ctenostoma (Myrmecilla) minusculum Naviaux 192 Ctenostoma (Myrmecilla) modicum Naviaux 192 Ctenostoma (Myrmecilla) obliquatum Chaudoir 194 Ctenostoma (Myrmecilla) oblitum Chaudoir 194 Ctenostoma (Myrmecilla) parvulum Naviaux 196 Ctenostoma (Myrmecilla) plicaticolle W. Horn 197 Ctenostoma (Myrmecilla) pygmaeum concolor Naviaux 197 Ctenostoma (Myrmecilla) pygmaeum pygmaeum (Lacordaire) 198 Ctenostoma (Myrmecilla) rivalieri Naviaux 199 Ctenostoma (Myrmecilla) schaumi pilosulum W. Horn 200 Ctenostoma (Myrmecilla) schaumi schaumi W. Horn 201 Ctenostoma (Myrmecilla) schmalzi paraguayense Brouerius van Nidek 201 Ctenostoma (Myrmecilla) schmalzi schmalzi W. Horn 201 Ctenostoma (Myrmecilla) unifasciatum affine W. Horn 204 Ctenostoma (Myrmecilla) unifasciatum unifasciatum Dejean 204

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Ctenostoma (Myrmecilla) vicinum Naviaux 205 Ctenostoma (Myrmecilla) zerchei Naviaux 206 Ctenostoma (Myrmecilla) zonata Chaudoir 206 Ctenostoma (Naviauxiana) aeneum Naviaux 173 Ctenostoma (Naviauxiana) arnaudi Naviaux 174 Ctenostoma (Naviauxiana) cayennense Naviaux 178 Ctenostoma (Naviauxiana) dormeri W. Horn 181 Ctenostoma (Naviauxiana) durantoni Naviaux 181 Ctenostoma (Naviauxiana) ebenina Bates 182 Ctenostoma (Naviauxiana) ecuadorense Naviaux 182 Ctenostoma (Naviauxiana) erwini Naviaux 182 Ctenostoma (Naviauxiana) insigne Chaudoir 188 Ctenostoma (Naviauxiana) longipalpe Naviaux 190 Ctenostoma (Naviauxiana) maculosum Naviaux 191 Ctenostoma (Naviauxiana) metallicum (Laporte de Castelnau) 192 Ctenostoma (Naviauxiana) nigrum lecourti Naviaux 193 Ctenostoma (Naviauxiana) nigrum nigrum Chaudoir 193 Ctenostoma (Naviauxiana) onorei colombiense Naviaux 195 Ctenostoma (Naviauxiana) onorei onorei Naviaux 195 Ctenostoma (Naviauxiana) ornatum ornatum Klug 195 Ctenostoma (Naviauxiana) ornatum werneri Naviaux 195 Ctenostoma (Naviauxiana) panamense Naviaux & Brzoska 196 Ctenostoma (Naviauxiana) pearsoni Naviaux 196 Ctenostoma (Naviauxiana) sallei Chaudoir 200 Ctenostoma (Naviauxiana) simile Naviaux 201 Ctenostoma (Naviauxiana) spinosum Naviaux 202 Ctenostoma (Naviauxiana) subtilesculptum W. Horn 202 Ctenostoma (Neoprocephalus) angustoobliquatum W. Horn 174 Ctenostoma (Neoprocephalus) brendelli Naviaux 176 Ctenostoma (Neoprocephalus) brunneum Naviaux 178 Ctenostoma (Neoprocephalus) cassolai Naviaux 178 Ctenostoma (Neoprocephalus) cylindratum Naviaux 180 Ctenostoma (Neoprocephalus) davidsoni Naviaux 180 Ctenostoma (Neoprocephalus) deuvei Naviaux 181 Ctenostoma (Neoprocephalus) germaini W. Horn 184 Ctenostoma (Neoprocephalus) guatemalense Brouerius van Nidek 185 Ctenostoma (Neoprocephalus) hovorei Naviaux & Brzoska 186 Ctenostoma (Neoprocephalus) ibidion ibidion Dohrn 186 Ctenostoma (Neoprocephalus) ibidion jolyi Naviaux 186 Ctenostoma (Neoprocephalus) immaculatum W. Horn 187 Ctenostoma (Neoprocephalus) intermedium Naviaux 188 Ctenostoma (Neoprocephalus) johnsoni Naviaux 189 Ctenostoma (Neoprocephalus) laeticolor Bates 189

Ctenostoma (Neoprocephalus) landolti Steinheil 190 Ctenostoma (Neoprocephalus) maculicorne (Chevrolat) 191 Ctenostoma (Neoprocephalus) nitidum Naviaux 194 Ctenostoma (Neoprocephalus) tumidum Naviaux 203 Ctenostoma (Neoprocephalus) turnbowi Naviaux 204 Ctenostoma (Neoprocephalus) vairai Cassola 205 Ctenostoma (Neoprocephalus) wappesi Naviaux 205 Ctenostoma (Paractenosoma) corculum Bates 179 Ctenostoma (Paractenosoma) parallelum Navaiux 196 Ctenostoma (Paractenosoma) simpliceps W. Horn 202 Ctenostoma (Salvioides) zikani W. Horn 206 Cychrini Laporte de Castelnau 137 Cychrus Fabricius 137 Cychrus hemphillii hemphillii G. Horn 138 Cychrus hemphillii rickseckeri LeConte 138 Cychrus tuberculatus T.W. Harris 138 L Leistus Frölich 22 Leistus (Leistus) ferrugineus (Linné) 22 Leistus (Neoleistus) ferruginosus von Mannerheim 23 Leistus (Neoleistus) longipennis Casey 23 Leistus (Neoleistus) madmeridianus Erwin 24 Loricera Latreille 69 Loricera aptena Ball & Erwin 70 Loricera decempunctata Eschscholtz 70 Loricera foveata LeConte 70 Loricera pilicornis congesta von Mannerheim 71 Loricera pilicornis pilicornis (Fabricius) 71 Loricera rotundicollis Chaudoir 72 Loricerini LeConte 69 N Nebria Latreille 24 Nebria (Boreonebria) crassicornis crassicornis Van Dyke 29 Nebria (Boreonebria) crassicornis intermedia Van Dyke 29 Nebria (Boreonebria) frigida R.F. Sahlberg 31 Nebria (Boreonebria) gouleti Kavanaugh 33 Nebria (Boreonebria) gyllenhali castanipes (Kirby) 34 Nebria (Boreonebria) gyllenhali lassenensis Kavanaugh 34 Nebria (Boreonebria) gyllenhali lindrothi Kavanaugh 35 Nebria (Boreonebria) hudsonica LeConte 35 Nebria (Boreonebria) lacustris bellorum Kavanaugh 37 Nebria (Boreonebria) lacustris lacustris Casey 38 Nebria (Boreonebria) nivalis gaspesiana Kavanaugh 41 Nebria (Boreonebria) nivalis nivalis (Paykull) 41 Nebria (Boreonebria) paradisi Darlington 43

INDEX

Nebria (Boreonebria) turmaduodecima Kavanaugh 48 Nebria (Catonebria) calva Kavanaugh 28 Nebria (Catonebria) carri Kavanaugh 28 Nebria (Catonebria) catenata Casey 28 Nebria (Catonebria) coloradensis Van Dyke 29 Nebria (Catonebria) gebleri albimontis Kavanaugh 32 Nebria (Catonebria) gebleri cascadensis Kavanaugh 32 Nebria (Catonebria) gebleri fragariae Kavanaugh 32 Nebria (Catonebria) gebleri gebleri Dejean 32 Nebria (Catonebria) gebleri rathvoni LeConte 33 Nebria (Catonebria) gebleri siskiyouensis Kavanaugh 33 Nebria (Catonebria) ingens ingens G. Horn 36 Nebria (Catonebria) ingens riversi Van Dyke 36 Nebria (Catonebria) kincaidi balli Kavanaugh 36 Nebria (Catonebria) kincaidi kincaidi Schwarz 37 Nebria (Catonebria) labontei Kavanaugh 37 Nebria (Catonebria) meanyi giulianii Kavanaugh 39 Nebria (Catonebria) meanyi lamarckensis Kavanaugh 40 Nebria (Catonebria) meanyi meanyi Van Dyke 40 Nebria (Catonebria) meanyi sylvatica Kavanaugh 40 Nebria (Catonebria) metallica Fischer von Waldheim 40 Nebria (Catonebria) ovipennis LeConte 42 Nebria (Catonebria) piperi Van Dyke 44 Nebria (Catonebria) piute piute Erwin & Ball 44 Nebria (Catonebria) piute sevieri Kavanaugh 44 Nebria (Catonebria) piute utahensis Kavanaugh 44 Nebria (Catonebria) purpurata LeConte 45 Nebria (Catonebria) schwarzi beverlianna Kavanaugh 46 Nebria (Catonebria) schwarzi schwarzi Van Dyke 46 Nebria (Catonebria) sierrablancae Kavanaugh 46 Nebria (Catonebria) spatulata sierrae Kavanaugh 47 Nebria (Catonebria) spatulata spatulata Van Dyke 47 Nebria (Catonebria) steensensis Kavanaugh 47 Nebria (Catonebria) trifaria pasquineli Kavanaugh 48 Nebria (Catonebria) trifaria trifaria LeConte 48 Nebria (Catonebria) vandykei vandykei Bänninger 49 Nebria (Catonebria) vandykei wyeast Kavanaugh 49 Nebria (Nebria) brevicollis (Fabricius) 27 Nebria (Reductonebria) acuta acuta Lindroth 24 Nebria (Reductonebria) acuta quileute Kavanaugh 25 Nebria (Reductonebria) acuta sonorae Kavanaugh 25 Nebria (Reductonebria) appalachia Darlington 25 Nebria (Reductonebria) arkansana arkansana Casey 26 Nebria (Reductonebria) arkansana edwardsi Kavanaugh 26 Nebria (Reductonebria) arkansana fragilis Casey 26 Nebria (Reductonebria) arkansana oowah Kavanaugh 27 Nebria (Reductonebria) charlottae Lindroth 28

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Nebria (Reductonebria) danmanni Kavanaugh 30 Nebria (Reductonebria) darlingtoni Kavanaugh 30 Nebria (Reductonebria) desolata Kavanaugh 30 Nebria (Reductonebria) diversa LeConte 30 Nebria (Reductonebria) eschscholtzii Ménétriés 31 Nebria (Reductonebria) gregaria Fischer von Waldheim 33 Nebria (Reductonebria) haida Kavanaugh 35 Nebria (Reductonebria) jeffreyi Kavanaugh 36 Nebria (Reductonebria) lituyae Kavanaugh 38 Nebria (Reductonebria) louiseae Kavanaugh 38 Nebria (Reductonebria) lyelli Van Dyke 39 Nebria (Reductonebria) mannerheimii Fischer von Waldheim 39 Nebria (Reductonebria) navajo Kavanaugh 41 Nebria (Reductonebria) obliqua chuskae Kavanaugh 42 Nebria (Reductonebria) obliqua obliqua LeConte 42 Nebria (Reductonebria) pallipes Say 43 Nebria (Reductonebria) sahlbergii modoc Kavanaugh 45 Nebria (Reductonebria) sahlbergii sahlbergii Fischer von Waldheim 45 Nebria (Reductonebria) sahlbergii triad Kavanaugh 46 Nebria (Reductonebria) suturalis LeConte 47 Nebria (Reductonebria) wallowae Kavanaugh 49 Nebria (Reductonebria) zioni oasis Kavanaugh 49 Nebria (Reductonebria) zioni zioni Van Dyke 50 Nebriini Castelnau de Laporte 22 Nippononebria Uéno 50 Nippononebria (Vancouveria) altisierrae (Kavanaugh) 50 Nippononebria (Vancouveria) campbelli (Kavanaugh) 51 Nippononebria (Vancouveria) virescens (G. Horn) 51 Notiokasiini Kavanaugh & Nègre 51 Notiokasis Kavanaugh & Nègre 51 Notiokasis chaudoiri Kavanaugh & Nègre 52 Notiophilini Mostschulsky 53 Notiophilus Dumeril 53 Notiophilus aeneus (Herbst) 54 Notiophilus aquaticus (Linné) 54 Notiophilus biguttatus (Fabricius) 55 Notiophilus borealis Harris 56 Notiophilus chihuahuae Casey 56 Notiophilus directus Casey 56 Notiophilus intermedius Lindroth 57 Notiophilus nemoralis Fall 57 Notiophilus nitens LeConte 57 Notiophilus novemstriatus LeConte 57 Notiophilus palustris (Duftschmid) 58 Notiophilus rufipes Curtis 59

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Notiophilus semiopacus Eschscholtz 59 Notiophilus semistriatus Say 59 Notiophilus sierranus Casey 60 Notiophilus simulator Fall 60 Notiophilus specularis Bates 60 Notiophilus sylvaticus Eschscholtz 61 O Omophron Latreille 63 Omophron americanum Dejean 64 Omophron baenningeri Dupuis 64 Omophron dentatum LeConte 64 Omophron dominicense Chaudoir 65 Omophron gilae LeConte 65 Omophron gratum Chaudoir 65 Omophron grossum Casey 66 Omophron labiatum (Fabricius) 66 Omophron mexicanum Dupuis 66 Omophron nitidum LeConte 66 Omophron obliteratum G. Horn 67 Omophron oblongiusculum Chevrolat 67 Omophron ovale G. Horn 67 Omophron robustum G. Horn 68 Omophron solidum Casey 68 Omophron sphaericum Chevrolat 68 Omophron tessellatum Say 69 Omophronini Bonelli 63 Opisthiini Dupuis 52 Opisthius Kirby 52 Opisthius richardsoni Kirby 53 P Pelophila Dejean 20 Pelophila borealis (Paykull) 20 Pelophila rudis (LeConte) 21 Pelophilini Kavanaugh 20 S Scaphinotus Dejean 1826 139 Scaphinotus (Brennus) bullatus Van Dyke 145 Scaphinotus (Brennus) cordatus (LeConte) 145 Scaphinotus (Brennus) crenatus (Motschulsky) 146 Scaphinotus (Brennus) cristatus (T.W. Harris) 146 Scaphinotus (Brennus) interruptus (Ménétriés) 152 Scaphinotus (Brennus) johnsoni Van Dyke 153 Scaphinotus (Brennus) marginatus (Fischer von Waldheim) 155

Scaphinotus (Brennus) obliquus (LeConte) 156 Scaphinotus (Brennus) oreophilus (Rivers) 156 Scaphinotus (Brennus) punctatus (LeConte) 159 Scaphinotus (Brennus) riversi (Roeschke) 161 Scaphinotus (Brennus) rugiceps incipiens (Casey) 161 Scaphinotus (Brennus) rugiceps rugiceps (G. Horn) 161 Scaphinotus (Brennus) striatopunctatus (Chaudoir) 163 Scaphinotus (Brennus) subtilis (Schaum) 163 Scaphinotus (Brennus) ventricosus (Dejean) 166 Scaphinotus (Maronetus) hubbardi (Schwarz) 150 Scaphinotus (Maronetus) imperfectus (G. Horn) 151 Scaphinotus (Maronetus) incompletus (Schwarz) 151 Scaphinotus (Maronetus) schwarzi (Beutenmüller) 162 Scaphinotus (Maronetus) tenuis (Casey) 164 Scaphinotus (Maronetus) unistriatus (Darlington) 165 Scaphinotus (Neocychrus) angulatus (T.W. Harris) 143 Scaphinotus (Neocychrus) behrensi (Roeschke) 144 Scaphinotus (Neocychrus) longiceps Van Dyke 154 Scaphinotus (Nomaretus) bilobus (Say) 144 Scaphinotus (Nomaretus) cavicollis (LeConte) 145 Scaphinotus (Nomaretus) debilis alpinus (Beutenmüller) 147 Scaphinotus (Nomaretus) debilis debilis (LeConte) 147 Scaphinotus (Nomaretus) fissicollis (LeConte) 149 Scaphinotus (Nomaretus) infletus Allen & Carlton 151 Scaphinotus (Nomaretus) liebecki Van Dyke 153 Scaphinotus (Pseudonomaretus) mannii Wickham 154 Scaphinotus (Pseudonomaretus) merkelii (G. Horn) 155 Scaphinotus (Pseudonomaretus) regularis (LeConte) 159 Scaphinotus (Pseudonomaretus) relictus (G. Horn) 160 Scaphinotus (Scaphinotus) elevatus coloradensis Van Dyke 147 Scaphinotus (Scaphinotus) elevatus elevatus (Fabricius) 148 Scaphinotus (Scaphinotus) elevatus flammeus Haldeman 148 Scaphinotus (Scaphinotus) elevatus lengi Van Dyke 148 Scaphinotus (Scaphinotus) elevatus neomexicanus Van Dyke 149 Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke 149 Scaphinotus (Scaphinotus) kelloggi (Dury) 153 Scaphinotus (Scaphinotus) macrogonus Bates 154 Scaphinotus (Scaphinotus) mexicanus Bates 156 Scaphinotus (Scaphinotus) parisianus Allen & Carlton 157 Scaphinotus (Scaphinotus) petersi biedermani Roeschke 157 Scaphinotus (Scaphinotus) petersi catalinae Van Dyke 157 Scaphinotus (Scaphinotus) petersi corvus (Fall) 158 Scaphinotus (Scaphinotus) petersi grahami Van Dyke 158 Scaphinotus (Scaphinotus) petersi kathleenae Ball 158 Scaphinotus (Scaphinotus) petersi petersi Roeschke 159 Scaphinotus (Scaphinotus) snowi parkeri Van Dyke 162

INDEX

Scaphinotus (Scaphinotus) snowi roeschkei Van Dyke 162 Scaphinotus (Scaphinotus) snowi snowi (LeConte) 163 Scaphinotus (Scaphinotus) unicolor (Fabricius) 164 Scaphinotus (Scaphinotus) vandykei Roeschke 165 Scaphinotus (Scaphinotus) viduus (Dejean) 166 Scaphinotus (Scaphinotus) webbi Bell 167 Scaphinotus (Steniridia) aeneicollis (Beutenmüller) 139 Scaphinotus (Steniridia) andrewsi amplicollis (Casey) 140 Scaphinotus (Steniridia) andrewsi andrewsi (T.W. Harris) 140 Scaphinotus (Steniridia) andrewsi barksdalei (Valentine) 140 Scaphinotus (Steniridia) andrewsi darlingtoni (Valentine) 141 Scaphinotus (Steniridia) andrewsi germari (Chaudoir) 141 Scaphinotus (Steniridia) andrewsi montanus (Valentine) 141 Scaphinotus (Steniridia) andrewsi mutabilis (Casey) 142 Scaphinotus (Steniridia) andrewsi nantahalae (Valentine) 142 Scaphinotus (Steniridia) andrewsi parvitarsalis (Valentine) 142 Scaphinotus (Steniridia) andrewsi saludae (Valentine) 143 Scaphinotus (Steniridia) andrewsi waldensius (Valentine) 143 Scaphinotus (Steniridia) guyotii (LeConte) 150 Scaphinotus (Steniridia) lodingi lodingi (Valentine) 154 Scaphinotus (Steniridia) lodingi obscurus (Valentine) 153 Scaphinotus (Steniridia) ridingsii intermedius Valentine 160 Scaphinotus (Steniridia) ridingsii monongahelae Leconte 160 Scaphinotus (Steniridia) ridingsii ridingsii (Bland) 160 Scaphinotus (Steniridia) tricarinatus (Casey) 164 Scaphinotus (Steniridia) violaceus carolinae (Valeneitne) 167

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Scaphinotus (Steniridia) violaceus violaceus (LeConte) 167 Scaphinotus (Stenocantharis) angusticollis (Fischer von Waldheim) 144 Scaphinotus (Stenocantharis) hatchi Beer 150 Scaphinotus (Stenocantharis) velutinus (Ménétriés) 165 Sphaeroderus Dejean 167 Sphaeroderus bicarinatus (LeConte) 168 Sphaeroderus canadensis canadensis Chaudoir 168 Sphaeroderus canadensis lengi Darlington 169 Sphaeroderus indianae (Blatchley) 169 Sphaeroderus nitidicollis brevoorti LeConte 169 Sphaeroderus nitidicollis nitidicollis Guérin-Méneville 170 Sphaeroderus nitidicollis schaumii Chaudoir 170 Sphaeroderus stenostomus lecontei Dejean 170 Sphaeroderus stenostomus stenostomus (Weber) 171 Systolosoma Solier 17 Systolosoma breve Solier 17 Systolosoma lateritium Nègre 18 Systolosomatini Erwin 17 T Trachypachini Thomson 18 Trachypachus Motschulsky 18 Trachypachus gibbsii LeConte 19 Trachypachus holmbergi von Mannerheim 19 Trachypachus slevini Van Dyke 19

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GENERAL BIBLIOGRAPHY FOR CARABOIDEA OF THE WESTERN HEMISPHERE

Plates

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Plate 1. Systolosoma Solier

TOP LEFT, extended focus image, adult of Systolosoma breve Solier from Osorno, Chile, ABL (adult length) 4.5 mm (ADP108775); TOP RIGHT, Systolosoma breve Solier, living beetle at P.N. Nahuelbuta, Chile, 1249 meters altitude; BOTTOM LEFT, typical landscape (Chile: Puente Blanco, 180 meters altitude) in which members of this genus dwell; BOTTOM RIGHT, habitat in which Systolosoma breve Solier lives. Living and habitat images credit: David R. Maddison, University of Arizona, Tucson. Landscape image credit: Allan Ashworth, North Dakota State University, Fargo.

Plate 2. Trachypachus Motschulsky

TOP LEFT, extended focus image, adult of Trachypachus holmbergi von Mannerheim from Twin Lakes, Alberta, Canada, ABL (adult length) 4.0 mm (ADP108776); TOP RIGHT, Trachypachus slevini Van Dyke, living beetle at Moolack Beach along the coast of Oregon; BOTTOM, typical landscape (Moolack Beach along the coast of Oregon) in which members of this genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson.

Plate 3. Pelophila Dejean

TOP, digital image, adult of Pelophila rudis (LeConte) from George Lake, Alberta, Canada, ABL (adult length) 13.0 mm (ADP108777); TOP RIGHT, Pelophila borealis (Paykull), living beetle at Castle Island, Quoile Pondage, County Down, N. Ireland, 5 meters altitude; BOTTOM, Buskin River, Kodiak Island, Alaska where adults of P. borealis Paykull are found under stones on rocky flats beside the river. Living and landscape image credits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 4. Leistus Frölich

TOP, extended focus image, adult of Leistus longipennis Casey from Prairie Creek, California, ABL (adult length) 9.5 mm (ADP108778); TOP RIGHT, Leistus rufomarginatus (Duftschmidt), living beetle at Castlewellan, County Down, N. Ireland, 150 meters altitude; BOTTOM, Lituya Bay, Glacier Bay National Park, Alaska where adults of L. ferruginosus Mannerheim are found under logs and on leaves and stems of Vaccinium sp. at night. Living and landscape image credits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 5. Nebria Latreille

TOP LEFT, extended focus image, adult of Nebria pallipes Say from Franklinville, Maryland, ABL (adult length) 10.0 mm (ADP108779); TOP RIGHT, Nebria metallica Fischer von Waldheim, living beetle at Montana Creek, NW of Juneau, Alaska; BOTTOM, typical landscape (Misery Bay Provincial Conservation area, Manitoulin Island, Ontario) in which members of this genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Henri Goulet, CNC, Ottawa, Ontario, respectively.

Plate 6. Nippononebria Uéno

TOP LEFT, extended focus image, adult of Nippononebria virescens (G. Horn) from Olympia, Washington, ABL (adult length) 10.0 mm (ADP108780); TOP RIGHT, Nippononebria virescens (G. Horn), living beetle at Stanley Park, Vancouver, British Columbia; Stanley Park, Vancouver, British Columbia; BOTTOM, typical habitat at Stanley Park where adults of N. virescens Horn are found hiding under flakes and in cracks on bare vertical wall of trail cut. Living and landscape images credit: David H. Kavanaugh, California Academy of Sciences, San Francisco.

Plate 7. Notiokasis Kavanaugh & Nègre

Extended focus image, adult of Notiokasis chaudoiri Kavanaugh & Nègre from Montevideo, Uruguay, ABL (adult length) 6.5 mm (in CAS). EFI credit: David H. Kavanaugh, California Academy of Sciences, San Francisco.

Plate 8. Opisthius Kirby

TOP LEFT, extended focus image, adult of Opisthius richardsoni Kirby from near Troutdal, Oregon, ABL (adult length) 10.0 mm (ADP108781); TOP RIGHT, Opisthius richardsoni Kirby, living beetle at Kootenay River near Bull River, BC, 750 meters altitude; BOTTOM, typical landscape (Ukak River, Valley of Ten Thousand Smokes, Katmai National Park, Alaska) where adults of O. richardsoni Kirby are found under stones at the edge of streams in this area. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and David H. Kavanaugh, California, Academy of Sciences, San Francisco, respectively.

Plate 9. Notiophilus Dumeril

TOP LEFT, extended focus image, adult of Notiophilus aeneus (Herbst) from near Largo, Maryland, ABL (adult length) 5.5 mm (ADP108782); TOP RIGHT, Notiophilus aeneus (Herbst), living beetle at Eno River State Park, NC, 135 meters altitude; BOTTOM, typical landscape (Plummers Island, Maryland, 80 meters altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Grace P. Servat, UNOP, respectively.

Plate 10. Cicindis Bruch

TOP LEFT, extended focus image of Cicindis horni Bruch, ABL (adult length) 9.5 mm (ADP108783), from Salinas Grande, Córdoba, Argentina; TOP RIGHT, Cicindis horni Bruch, living beetles, swarming at the Salinas Grande, Córdoba, Argentina; BOTTOM, actual landscape (Salinas Grande, Córdoba, Argentina ) in which members of this species dwell. Living and landscape images credit: Valeria Aschero, CRICyT, Mendoza, Argentina.

Plate 11. Omophron Latreille

TOP LEFT, extended focus image of Omophron americanum Dejean from Plummers Island, Maryland, ABL (adult length) 6.0 mm (ADP034029); TOP RIGHT, Omophron tessellatum Say, living beetle at Boone River near Webster City, IA, 305 meters altitude; BOTTOM, typical landscape (sandy beach, Lake Huron) in which members of this genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Henri Goulet, CNC, Ottawa, Ontario, respectively.

Plate 12. Loricera Latreille

TOP, extended focus image of Loricera foveata LeConte, from Tacoma, Washington, ABL (adult length) 7.5 mm (ADP108784); TOP RIGHT, Loricera pilicornis (Fabricius), living beetle at Newtownbreda, Belfast, N. Ireland, 50 meters altitude; BOTTOM, typical landscape (12.9 km S of Dixon, Solano County, California, 30 meters altitude) where adults of L. foveata LeConte are found under debris in this grassy habitat. Living and landscape image credits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 13. Callisthenes Fischer von Waldheim

TOP, digital image of Callisthenes affinis (Chaudoir), from near Garfield, Oklahoma, ABL (adult length) 24.0 mm (ADP108787); BOTTOM, typical landscape (Plummers Island, Maryland, 80 meters altitude) in which members of the genus dwell. Landscape image credit: Grace P. Servat, UNOP.

Plate 14. Calosoma (Callitropa Motschulsky)

TOP, digital image, adult of Calosoma externum (Say), from Washington, DC, ABL (adult length) 25.0 mm (ADP108785); BOTTOM, typical landscape (Plummers Island, Maryland, 80 meters altitude) in which members of the genus dwell. Landscape image credit: Grace P. Servat, UNOP

Plate 15. Calopachys Haury

TOP, digital image, adult of Calopachys viridissimus Haury, from 10 km NW Oaxaca, Oaxaca, México, ABL (adult length) 21.0 mm (ADP108786); BOTTOM, typical landscape (N of Oaxaca, México) in which members of the genus dwell. Landscape image credit: Christopher J. Earle, Gymnosperm Database (http://www.conifers.org).

Plate 16. Calosoma Weber

TOP LEFT, digital image, adult of Calosoma scrutator (Fabricius) from near Snow Hill, Maryland, ABL (adult length) 28.0 mm (ADP108785); TOP RIGHT, Calosoma scrutator (Fabricius), living beetle at Walker Canyon, AZ, 1215 meters altitude; BOTTOM, typical landscape (Plummers Island, Maryland, 80 meters altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Grace P. Servat, UNOP, respectively.

Plate 17. Carabus Linné

TOP LEFT, digital image, adult of Carabus serratus Say from Plummers Island, Maryland, ABL (adult length) 19.0 mm (ADP020031); TOP RIGHT, Carabus vietinghoffi Adams, living beetle at Pelley Crossing, Pelley River, Yukon; BOTTOM, St. Paul Island, Pribilof Islands, Alaska, where adults of C. truncaticollis Eschscholtz are found on coastal tundra under debris. Living and landscape images credit: Henri Goulet, CNC, Ottawa, Ontario and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 18. Ceroglossus Solier

TOP LEFT, digital image, adult of Ceroglossus magellanicus araucanus Kraatz-Koschlau, from Araucania, Chile, ABL (adult length) 25.0 mm (ADP108789); TOP RIGHT, Ceroglossus ochensii (Germain) living beetle at Chiloé, above Rio Cipresal, 195 meters altitude; BOTTOM, typical landscape (Mallines, Parque Nacional Puyehue at 710m altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Allan Ashworth, North Dakota State University, Fargo, respectively.

Plate 19. Cychrus Fabricius

TOP, digital image, adult of Cychrus tuberculatus T.W. Harris, from Florence, Oregon, ABL (adult length) 21.0 mm (ADP108791); TOP RIGHT, Cychrus caraboides (Linné), living beetle at Hillsborough, County Down, N. Ireland, 110 meters altitude; BOTTOM, Lepas Bay area, Graham, Island, Queen Charlotte Islands, British Columbia where adults of C. tuberculatus Harris are found walking on the forest floor at night. Living and landscape image credits: Roy Anderson, Belfast, Northern Ireland, U.K. and David H. Kavanaugh, California Academy of Sciences, San Francisco, respectively.

Plate 20. Scaphinotus Dejean

TOP LEFT, digital image, adult of Scaphinotus ridingsii ridingsii (Bland), from Plummers Island, Maryland, ABL (adult length) 16.0 mm (ADP020069); TOP RIGHT, Scaphinotus marginatus (Fischer von Waldheim), living beetle at Eagle River Trail, N. of Juneau, AK, 6 meters altitude; BOTTOM, typical landscape (Plummers Island, Maryland, 80 meters altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson and Grace P. Servat, UNOP, respectively.

Plate 21. Sphaeroderus Dejean

TOP LEFT, digital image, adult of Sphaeroderus bicarinatus (LeConte) from Yellow Face, North Carolina, ABL (adult length) 16.0 mm (ADP0108790); TOP RIGHT, Sphaeroderus stenostomus stenostomus (Weber), living beetle at Eno River State Park, NC, 135 meters altitude; BOTTOM, typical landscape (Eno River, State Park, NC, 135 meters altitude) in which members of the genus dwell. Living and landscape images credit: David R. Maddison, University of Arizona, Tucson.

Plate 22. Ctenostoma Klug

TOP, digital image, adult of Ctenostoma erwini Naviaux, ABL (adult length) 13.5 mm (ADP000530), from Barro Colorado Island, Panamá; BOTTOM, typical suspended dry leaf habitat (Tiputini Biodiversity Station, Ecuador) in which members of the genus dwell. Landscape image credit: Terry L. Erwin.